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Volume 36 1982 Number |] 


ISSN 0024-0966 


_ JOURNAL 


LEPIDOPTERISTS’ SOCIETY 


Published quarterly by THE LEPIDOPTERISTS’ SOCIETY 


Publié par LA SOCIETE DES LEPIDOPTERISTES 
Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN 
Publicado por LA SOCIEDAD DE I.OS LEPIDOPTERISTAS 


16 June 1982 


THE LEPIDOPTERISTS’ SOCIETY 


EXECUTIVE COUNCIL 


LINCOLN P. BROWER, President C. R. BEUTELSPACHER B.., 
MARIA ETCHEVERRY, Ist Vice President Immediate Past President 

G. L. MULLER, Vice President JULIAN P. DONAHUE, Secretary 
R. H. CARCASSON, Vice President RONALD LEUSCHNER, Treasurer 


Members at large: 


M. DEANE BOWERS R. L. LANGSTON K. S. BROWN, JR. 
E. R. HODGES R. M. PYLE T. C. EMMEL 
W. D. WINTER A. M. SHAPIRO 


The object of the Lepidopterists’ Society, which was formed in May, 1947 and for- 
mally constituted in December, 1950, is “to promote the science of lepidopterology in 
all its branches, ....to issue a periodical and other publications on Lepidoptera, to 
facilitate the exchange of specimens and ideas by both the professional worker and the 
amateur in the field; to secure cooperation in all measures” directed towards these 
aims. 

Membership in the Society is open to all persons interested in the study of Lepi- 
doptera. All members receive the Journal and the News of the Lepidopterists Society. 
Institutions may subscribe to the Journal but may not become members. Prospective 
members should send to the Treasurer full dues for the current year, together with 
their full name, address, and special lepidopterological interests. In alternate years a 
list of members of the Society is issued, with addresses and special interests. There 
are four numbers in each volume of the Journal, scheduled for February, May, August 
and November, and six numbers of the News each year. 


Active members—annual dues $18.00 
Student members—annual dues $13.00 
Sustaining members—annual dues $25.00 
Life members—single sum $250.00 
Institutional subscriptions—annual $25.00 


Send remittances, payable to The Lepidopterists’ Society, and address changes to: 
Ronald Leuschner, 1900 John St., Manhattan Beach, California 90266 U.S.A. 


Back issues of the Journal of the Lepidopterists’ Society, the Commemorative Vol- 
ume, and recent issues of the NEWS are available from the Treasurer. The Journal is 
$13 per volume, the Commemorative Volume, $6; and the NEWS, $.25 per issue. 


Order: Mail to Ronald Leuschner, 1900 John St., Manhattan Beach, California 90266 
U.S.A. 


Journal of the Lepidopterists’ Society (ISSN 0024-0966) is published quarterly 
by the Lepidopterists’ Society, a non-profit, scientific organization. The known office 
of publication is 1041 New Hampshire St., Lawrence, Kansas 66044. Second class 
postage paid at Lawrence, Kansas, U.S.A. 66044. 


Cover illustration: Mature larva of Eumorpha fasciata Sulzer (Sphingidae) feeding 
on Ludwigia sp. (Jussiaea) in southern Florida, where this hawk moth is generally 
found throughout the year. Original drawing by Mr. John V. Calhoun, 382 Tradewind 
Ct., Westerville, Ohio 43081, USA. 


JOURNAL OF 


Tue LEPIDOPTERISTS’ SOCIETY 


Volume 36 1982 Number 1 


Journal of the Lepidopterists’ Society 
Sol) 1982. 1—17 


THE BUTTERFLY FAUNA OF BARTON CREEK CANYON ON 
THE BALCONES FAULT ZONE, AUSTIN, TEXAS, 
AND A REGIONAL LIST 


CHRISTOPHER J. DURDEN 


Curator of Geology and Entomology, Texas Memorial Museum, 
2400 Trinity Street, Austin, Texas 78705 


ABSTRACT. Diversity of substrate, topography and water supply, and climate and 
vegetation account for the occurrence of 74% of the regional fauna along a 1.1 km 
stretch of Barton Creek. Monthly mean weather records for 40 years are analyzed on 
a bioclimagram and the modes matched with habitat types. 172 species are listed for 
the ten counties around Austin, and range, habitat, abundance, and residency are in- 
dicated for the 127 found at the study site. Faunal history is interpreted as a sequence 
of colonization events followed by episodes of habitat restriction. These are related to 
regional patterns of climatic change inferred from the 22,000-year paleontologic record 
of sphagnum bogs 70 km to the east and southeast. No species listed is endangered by 
collecting. Several restricted species of special habitats are threatened by potentially 
careless land use. 


Barton Creek rises in Hays Co. on the Edwards Plateau and flows 
east into Travis Co. to enter the Colorado River at Austin below the 
Balcones Fault Zone. Most of the basin contains juniper and live-oak 
woodland, with cedar-elm more frequent towards the east. Post oak 
savanna occurs on higher hills near the headwaters, including Shingle 
Hills at 460 m. Madrone (Arbutus xalapensis H.B.K.), associated with 
sacahuista-grassland (Nolina lindheimeriana (Scheele) Wats. and N. 
texana Wats.) is found on the divides. Dwarf palmetto (Sabal minor 
(Jacq.) Pers.) and bald cypress (Taxodium distichum (L.) Rich.) are 
found in the deepest canyon in the middle part of the drainage (Cor- 
rell & Johnston, 1970). In terms of biotic diversity, the most interest- 
ing area is the lowest portion of the canyon where it crosses the spring 
zone of the Balcones Faults. Here, since 1971, have been taken 127 
species and an additional 10 subspecies in overlap. This is 74% of the 
total butterfly fauna known from the 10-county area around Austin 
(Llano, Burnet, Blanco, Hays, Travis, Williamson, Lee, Bastrop, Cald- 


2 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


well, and Fayette) or within approximately 90 km (Fig. 1). Factors 
accounting for this biotic diversity include: features of substrate; to- 
pography and water supply; and climate and vegetation. 

Substrate. Bedrock is a hard gray Cretaceous limestone (Rodda et 
al., 1970); the upper three members of the Edwards Formation, with 
the softer Georgetown Limestone capping hills north and south of the 
study area. The uppermost member of the Edwards is cavernous with 
rapid sink-in of rainfall, leading to development of rather xeric mi- 
crohabitats. Next below is the major cliff-forming member. Residual 
soils on these limestones are dark-brown and gray-brown granular 
with limestone fragments and are of the Tarrant soil group (Werchan 
et al., 1974). On steeper slopes occurs colluvium of broken limestone 
of the Brackett soil group. At approximately 170 m there are deposits 
of the Capitol Terrace of reddish-brown chert pebbles and clay on 
which are developed soils of the Speck group. This terrace was de- 
posited during or before the Yarmouth Interglacial, two to three 
hundred thousand years ago, when the gradient of the Colorado River 
and its tributaries was raised in adjustment with the interglacial rise 
in sea level. At approximately 150 m there is the Sixth Street Terrace 
of brown-red silty clay on which is developed soils of the Altoga 
group. This terrace deposit accumulated during the Sangamon Inter- 
glacial between 125,000 and 40,000 years ago. Insect microfossils in 
this deposit indicate a climate as warm as, or warmer than, present. 
At 140 m there is the First Street Valley Fill of brown fine sandy loam 
on which is formed soil of the Hardeman group. This deposit, which 
contains extinct horse and mastodon remains, was formed during the 
major pluvial stage in the late Wisconsin Glacial between 15,000 and 
9000 years ago. Insect microfossils in this deposit indicate a climate 
cooler and more moist than present. Four lower levels of valley fill 
occur to the east of the study site, deposited during later Wisconsin 
and post-glacial pluvial episodes. The present bed load of Barton 
Creek consists of gray silt with limestone boulders, which are accu- 
mulating since the damming and modification of the creek channel at 
Barton Springs swimming pool. 

Topography, land use and water supply. The collecting site 
extends from 1.2 km above the mouth of Barton Creek at the upper 
end of the swimming pool at 144 m elevation to 2.3 km SSW of the 
mouth at 157 m elevation above the rapids above Campbell’s Hole. 
In the channel there is a small perennial spring at 145 m and two 
usually perennial pools at 147 m, the uppermost of which has been 
known for 100 years as Campbell’s Hole. There is a sloping meadow 
at 150 to 155 m in the northeastern third of the area, upstream from 
which the canyon narrows. The top of the bluffs stands at approxi- 


VOLUME 36, NUMBER 1 3 


of WEST 
vA AUSTIN 
LE 


4 


LG 
-? CR: 


30° 20' N 

Fic. 1. Location of study site, with counties in the Austin area: 1. Llano, 2. Burmet, 
3. Blanco, 4. Hays, 5. Travis, 6. Williamson, 7. Lee, 8. Bastrop, 9. Caldwell, 10. Fayette; 
and substrate: A. Llano Uplift (Precambrian shield and Paleozoic limestones), B. Lower 
Cretaceous limestone plateau and canyons, C. Balcones Fault Zone (canyons and 
springs), D. Upper Cretaceous limestones (with black rendzina soils and prairie), E. 
Paleogene sandstones and shales (with barren sands, podsols and peat bogs), F. Neo- 
gene sands and clays (with coastal plain prairie). 


97°47-5' W 
97°45' W 


mately 185 m. The present boundary of Zilker City Park protects the 
north canyon wall almost to the top, but the south canyon wall lies 
outside the park, where it is not developed because a city regulation 
prohibits building on floodplains. Development in the form of apart- 
ment complexes and single family houses has, in the past 5 years, 
used up almost all land bordering the collecting site. In the early 
1960’s a major sewer line was laid along the northem border of the 
creek channel. Construction of this line cracked limestone ledges 
which formerly held ephemeral pools, the habitat of an endemic crus- 
tacean, Eulimnadia antlei Mackin, known elsewhere only from Okla- 


4 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Fic. 2. Bioclimagram for Austin, Texas, based on monthly means of temperature 
and rainfall from 1931 to 1970, summarized by point density contouring to locate 
modes. Apparent rainfall is monthly rainfall x12. 


homa. Published plans for future development of the study area in- 
clude a “Barton Creek Hike and Bikeway’ trail over the sewer line 
and wide enough for a standard patrol car to extend from Barton 
Springs to a major highway 3.5 km to the southwest. 

Climate and vegetation. Monthly rainfall and temperature records 
for Austin, covering the years 1931 to 1970 were plotted as points on 
a diagram similar to that of Holdridge et al. (1971). Points were standard- 
ized as if for yearly averages and then point density was contoured to 
locate the positions of climatic modes found throughout the year (Fig. 
2). The interpretive overlay of Holdridge was originally calibrated 
empirically for equatorial climates of small annual amplitude. A new 
overlay (Fig. 3) has been calibrated empirically for strongly seasonal 
climates, using as labels the habitats selectively favored under each 
condition. In order of decreasing frequency of occurrence, our sum- 
mer (hotter than 16°C, 63°F) bioclimates are for dry forest (sabinal), 
warm desert, warm arid forest (encinal), with minor modes at warm 
thorn scrub, warm thorn woodland (including chaparral), warm mesic 
forest (2 modes), and warm moist forest. Our winter bioclimates are 
moderate park woodland (2 modes) and moderate groved meadow. 
Because the annual bioclimate is an oscillation between modes, now 
one, then another, community best adapted for particular modes is at 
a selective advantage. The cumulative result of such historical selec- 
tion is a mosaic or patchy environment in which diverse communities 
coexist side by side in sharp discontinuity with boundaries main- 
tained by small, but decisive, edaphic and topographic differences. 
Winter temperatures run some 1.5°C higher, and in summer, some 
2..5°C lower in the canyons than the published records from the prairie 
site at the airport (U.S. Dept. Commerce, NOAA-EDS, 1970). 

On Barton Creek there is sabinal or Juniperus ashei Buchh. wood- 


VOLUME 36, NUMBER | 5 


Sows saline abet 


RAINFALL DESERT MEADOW BOG 
TUNDRA TUNDRA TUNDRA 
TEMPERATURE —, 


TAIGA MUSKEG 
COLD 
STEPPE MOOR 
HEATH MOSS FOREST 
GROVED | PARK 
PRAIRIE 
MEADOW | WOODLAND 


DRY 
WOODLAND 


Ve 


Fic. 3. Key diagram showing optimal community types for various temperatures 
and apparent rainfall figures based on data from a large number of North American 
stations. 


land on the divides. Oak (Quercus fusiformis Small) and elm (Ulmus 
crassifolia Nutt.) woodland occurs on deeper‘dry soils. Oak (Q. sin- 
uata Walt.) and elm (U. americana L.) woodland occurs on mesic 
colluvial pockets. Oak (Q. texana Buckl.), redbud (Cercis canadensis 
L.), red buckeye (Aesculus pavia L.), hop tree (Ptelea trifoliata L.), 
and monilla (Ungnadia speciosa Endl.) woodland occurs on steeper 
shade slopes. Pecan (Carya illinoinensis (Wang.) K. Koch) and ash 
(Fraxinus americana L.) woodland occurs on alluvial soil. Cotton- 
wood (Populus deltoides Marsh), willow (Salix nigra Marsh), plane 
tree (Platanus occidentalis L.), and indigo bush (Amorpha fruticosa 
L.) occur along the stream channel. On sunslopes there is a chaparral 
of evergreen sumac (Rhus virens Gray), ebony (Diospyros texana 
Scheele), stretch berry (Forestiera pubescens Nutt.), oreja de raton 
(Bernardia myricaefolia (Scheele) Wats.), christmas bush (Eupato- 
rium havanense H.B.K.), china (Sapindus saponaria L.), lotebush 
(Ziziphus obtusifolia (T. & G.) Gray), cenizo (Leucophyllum frutes- 
cens (Berl.) I. M. Johnst.), granjeno (Celtis pallida Torr.), mescal bean 
(Sophora secundiflora (Ort.) DC.), kidney wood (Eysenhardtia texana 
Scheele), catclaw (Acacia roemeriana Scheele) and many other 
shrubs. The grassland of cleared areas is richly herbaceous and is 
returning rapidly to shrubland under pioneering jara dulce (Baccharis 
neglecta Britt.), mesquite (Prosopis glandulosa Torr.), huisache (Aca- 
cia smallii Isely), and retama (Parkinsonia aculeata L.), and to savan- 
na with coma (Bumelia lanuginosa (Michx.) Pers.) and hackberry 
(Celtis laevigata Willd. and C. lindheimeri Engelm.). The herbaceous 
and small shrub flora is very rich, particularly on the limestone cliffs 


6 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY — 


where nectar sources are found in all seasons. During moist periods 
there is a proliferation of herbs and vines on the woodland floor, 
including suitable larval foodplants for many of our periodic tropical 
colonists. In cedar thickets in autumn, drying plants of Heliotropium 
tenellum (Nutt.) Torr. are attractants for Danaus spp. 

Most years have minimum temperature no lower than —5°C (23°F) 
during the second week of January. New growth of chamaephytes — 
begins in February. The mean last frost date is 3 March with rare 
chance of near frost as late as mid-April. The first rainy season peaks 
in May and is separated in most years by a more or less severe dry 
season in June, July, and August from the second rainy season in 
September. Rarely the first frost occurs as early as late October or as 
late as January, with the mean first frost date on 28 November. A few 
butterfly species are adult only during, just before, or just after the 
first rainy season. These are Erynnis juvenalis, Megathymus colora- 
densis kendalli, Satyrium calanus falacer, Callophrys solatus, Fixsen- 
ia ontario autolycus, Megisto cymela, and Anthocharis midea an- 
nickae, and all are single brooded. Other numbered species on the 
list are two or more brooded per year and may be found during or 
following both rainy seasons, or all year long. Periodic tropical colo- 
nists are usually not found before late August in normal years. In 
years of moist spring, when there is abundant rainfall in the Rio 
Grande Valley between Sabinas Hidalgo and San Antonio, they may 
reach our area in May or June. In rare years of exceptionally mild 
winter, some of these tropical colonists may survive the cold. Neck 
(1978) has given an introduction to the periodicity of this phenome- 
non. Empirically, the greatest diversity or number of synchronic 
species occurs in mid-October, and the greatest population or number 
of individual butterflies on the wing occurs in late September. There 
is a smaller diversity peak in June and population peak in May. Di- 
verse collecting often persists well into November and occasionally 
into December. Taxa of temperate distribution enter hibernation dur- 
ing October, possibly in response to change in day-length. This stops 
the emergence of new adults of these species, freeing niche space 
and nectar sources for the periodic tropical colonists. From mid-Oc- 
tober through December the greater part of the adult population is 
composed of species with predominantly tropical distribution, a few 
endemic taxa, and a few adult hibernators. One species of the saca- 
huista-grassland association, not yet known from the study site but 
occurring on the left divide 11 km to the northwest, is Hesperia wood- 
gatei. This species has a single autumn brood flying from mid-October 
into early November. The 27 species indicated (*) may be found dur- 


VOLUME 36, NUMBER 1 Fi 


ing warm weather in mid-winter. In dry years, more species can be 
found in January than in July. 

Biogeographic notes. The subspecies is chosen as the working 
taxon for biogeographic analysis because it is the primary occupant 
of the niche (Durden, 1969). Different subspecies have slightly dif- 
ferent habitat requirements and accordingly their ranges are deter- 
mined by the distribution of such habitats. At this locality there are 
6 pairs and 2 triplets of syntopic subspecies. These are, however, 
rarely synchronic or competitive. They have different overall ranges, 
different habitat requirements, and are only found together during 
episodes of mutual dispersal. 

By habitat preference our butterflies are 53% of woodlands, 26% of 
desert and grassland, 12% of brushland, and 9% of arid woodland and 
thorn scrub. Geographically they are 39% in midrange, 41% at or near 
the northern boundary, 18% near the eastern boundary, 17% at the 
southern or southwestern boundary, and 7% at the western boundary. 
Range groups, position within range, frequency, and residency status 
are indicated in the list by the following symbols: 

Range groups. A—wide-ranged in dry disturbed sites (9) in mid- 
range). B—Great Plains (8 in midrange, 4 at southern edge, 4 at east- 
ern edge). C—warm temperate, subtropical and tropical in dry open 
sites (9 in midrange, 2 at eastern edge). D—subtropical arid thorn 
forest (1 in midrange, 3 at eastern edge, 3 at northern edge). E— 
eastern subtropical arid thorn scrub (2 in midrange, 2 at northern 
edge). F—Tamaulipan arid woodland (2 at northern edge). G—broad- 
ranged temperate and tropical woodland (1 in midrange). H—eastern 
deciduous forest and tropical woodland (1 at western edge). I—Gulf 
Coast and tropical woodland (9 in midrange). J—subtropical and 
southern Great Plains brushland (3 in midrange, | at eastern edge). 
K—easterm deciduous and subtropical montane woodland (2 in mid- 
range). L—eastern deciduous woodland (5 in midrange, 5 at south- 
western edge). M—Appalachian and Mississippi Basin woodland (4 
at southern edge, 4 at southwestern edge). N—Gulf Coast woodland 
(7 at western edge). O—westerm deciduous and subtropical montane 
woodland (6 at eastern edge). P—endemic taxa of the Sierra Madre 
Oriental and Balcones Escarpment (4 at northern edge). Q—endemic 
taxa of central Texas and northern Coahuila (1 in midrange, 2 at east- 
ern edge, 4 at northern edge). R—tropical woodland (4 in midrange, 
11 within northern part of range, 15 at northern edge). 

Position within range. n—at northern edge of range (30). nm—in 
the northern part of the range, rarely straying farther (11). m—in mid- 
range (54). e—at eastern edge of range (18). w—at western edge of 


8 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


range (8). s—at southern edge of range (8). sw—at southwestern edge 
of range (9). 

Frequency. a—abundant, but may be periodically scarce, although 
frequently occurring in such abundance that a series may be taken in 
a day (84). u—uncommon, few specimens have been taken, several 
years are required to accumulate a series (27). s—scarce, one to five 
individuals have been taken in 10 years (26). 

Residency status. p—periodic, strays appearing in our region after 
a moist spring or after dispersal by hurricane, usually raising one or 
more broods or persisting up to 9 years before extermination by severe 
winter freeze or by summer drought (46). r—permanent residents, 
known to have survived extreme drought or winter freeze of 25 to 100 
year severity. 


LIST OF THE BUTTERFLIES FOUND ON BARTON CREEK IN ZILKER 
PARK, AUSTIN, TEXAS, WITH ADDITIONAL SPECIES FOUND IN THE 
ADJACENT TEN COUNTIES 


(Dates given for scarce species only; author was collector unless 
noted otherwise; footnotes 1-28 follow the List.) 


PYRGINAE (HESPERIIDAE) 


1. Epargyreus clarus (Cramer, 1775) subsp., O-e-a-r. 
2. Chioides albofasciatus (Hewitson, 1867), R-n-s-p (14:x:72). 
3. C. zilpa zilpa (Butler, 1874), R-n-s-p (29:ix:76). . 

Urbanus proteus (Linné, 1758), Bastrop Co. (Heiligbrodt, 1870's). 

4. U. dorantes rauterbergi (Skinner, 1895), R-nm-s-p (1:x:72). 

Achalarus coyote (Skinner, 1892), Travis Co. (Bull Cr.). 

5. Thorybes pylades (Scudder, 1870), K-m-u-r. 
T. confusis Bell, 1922, Bastrop Co., Travis Co. (Bull Cr.). 
T. bathyllus (Abbot in Smith, 1797), Bastrop Co. 
Cogia outis (Skinner, 1894), Travis Co. (Bull Cr.). 
6. Bolla clytius (Godman & Salvin, 1897), R-n-s-p (2:x:77). 
7. Staphylus hayhurstii (Edwards, 1870), L-m-u-r. 
8. Systasea pulverulenta (Felder, 1869), E-n-a-r. 
9. Achlyodes mithridates tamenund (Edwards, 1871), R-nm-u-p.! 
Gesta invisus llano (Dodge, 1903), Llano, Travis, Bastrop Co’s.? 
10. Erynnis horatius (Scudder & Burgess, 1870), L-m-a-r. 
11. E. meridianus Bell, 1927, B-s-s-r (30:ix:76). 
12. E. juvenalis juvenalis (Fabricius, 1793), M-s-u-r. 

E. martialis (Scudder, 1869), Bastrop Co. (Sandy Cr.). 
13. E. funeralis (Scudder & Burgess, 1870), C-m-a-r. 

E. zarucco (Lucas, 1857), Travis Co. (Bull Cr.). 

E. baptisiae (Forbes, 1936), Travis Co. (Bull Cr.). 

E. burgessi (Skinner, 1914), Blanco Co. (Round Mt.; F. G. Schaupp). 
14a. *Syrichtus (Tuttia) communis communis (Grote, 1872), C-m-a-r.? 
14b. S. (T.) communis albescens (Plotz, 1884), D-n-a-p.* 

S. (T.) oileus (Linné, 1767), Travis Co. (Bull Cr.). 

15. S. (T.) philetas (Edwards, 1881), E-m-u-r.® 
16. Heliopetes laviana (Hewitson, 1868), R-n-s-p (13:x:71, 9:xi:72). 
17. Celotes nessus (Edwards, 1877), E-m-a-r. 


VOLUME 36, NUMBER | fe) 


18. 


Pholisora catullus (Fabricius, 1793), M-s-a-r. 
P. mejicana (Reakirt, 1866), Travis Co. (Bull Cr.). 


HESPERIINAE (HESPERIIDAE) 


19: 


Megathymus coloradensis kendalli Freeman, 1965, Q-e-u-r.® 

M. coloradensis reinthali Freeman, 1963, Caldwell, Williamson, Lee Co’s. 
Ancyloxipha numitor (Fabricius, 1793), Travis, Lee Co’s. 

A. arene (Edwards, 1871), Llano Co. (Enchanted Rock). 


. Copaeodes aurantiaca (Hewitson, 1868), C-e-a-r. 
. *C. minima (Edwards, 1870), L-m-a-r (with the genitalically distinct winter f. 


rayata Barnes & McDunnough, 1913). 


. *Atalopedes campestris (Boisduval, 1852), L-m-a-r (with a dark winter f.). 
. Hesperia viridis (Edwards, 1883), B-e-u-r. 


H. woodgatei (Williams, 1914), Travis Co. (Fourpoints & U. Barton Cr.). 
H. licinus (Edwards, 1871), Bastrop Co. (Heiligbrodt Cln.).’ 

H. meskei (Edwards, 1877), Bastrop Co. (Heiligbrodt Cln.). 

H. attalus (Edwards, 1871), Travis Co. (U. Shoal Cr.). 


. Polites (Polites) vibex brettoides (Edwards, 1883), L-m-u-r. 


P. (P.) themistocles (Latreille, 1824) subsp., Bastrop Co. (Paige). 


. P. (Wallengrenia) otho otho (Abbot in Smith, 1797), L-m-a-r. 


Poanes (Poanes) viator viator (Edwards, 1865), Lee Co. (Patschke Bog). 


. Hylephila phyleus (Drury, 1773), L-m-a-r. 


Atrytone (Atrytone) arogos iowa (Scudder, 1868), Travis Co.’ 
A. (A.) arogos arogos (Boisduval & LeConte, 1833), Fayette Co. (Swissalp). 


. A. (A.) delaware lagus (Edwards, 1881), B-s-u-r. 


A. (A.) mazai Freeman, 1969, Travis Co. (U. Shoal Cr.). 


. Euphyes vestris osyka (Edwards, 1867), L-m-a-r. 
. Amblyscirtes nysa Edwards, 1877, B-e-u-r. 


A. aenus aenus Edwards, 1878, Travis Co. (Bull Cr.). 
A. erna Freeman, 1943, Travis Co. (Bull Cr.). 


. A. eos (Edwards, 1871), B-e-u-r. 
. A. celia Skinner, 1895, F-n-a-r. 


A. alternata (Grote & Robinson, 1867), Bastrop, Lee Co’s. 


. Lerodea eufala (Edwards, 1869), L-m-a-r. 

. Calpodes ethlius (Stoll, 1782), R-m-u-p. 

. Panoquina ocola (Edwards, 1863), L-m-a-r. 

. Nastra julia (Freeman, 1945), Q-m-a-r. 

. Lerema accius (Abbot in Smith, 1797), L-m-a-r. 


RIODININAE (LYCAENIDAE) 


37. 
38. 
39. 


40. 


Calephelis nemesis australis (Edwards, 1877), Q-n-a-r (with a winter form ap- 
proaching typical nemesis).° 

C. perditalis (Barnes & McDunnough, 1918), P-n-u-r (with darker angulate- 
winged winter form). 

C. guadeloupe (Strecker, 1878), Q-n-a-r (with summer f. rawsoni McAlpine, 
1939).1° 

C. sinaloensis nuevoleon McAlpine, 1971, P-n-s-r (16, 21, 28:x:71; 10:ix:72). 
Apodemia mormo mejicanus (Behr, 1865), Bastrop, Travis Co's." 


LIPHYRINAE (LYCAENIDAE) 


41. 


Feniseca tarquinius (Fabricius, 1793), M-sw-s-r (24:ix:79). 


LYCAENINAE (LYCAENIDAE) 


Chlorostrymon simaethis sarita (Skinner, 1895), Williamson Co. (Salado Cr.). 
Phaeostrymon alcestis alcestis (Edwards, 1871), Williamson, Travis Co's. 


42. Satyrium calanus falacer (Godart, 1824), M-s-u-r. 


10 


43. 


44. 


61. 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Calycopis isobeon (Butler & Druce, 1869), J-m-a-r (with a dimorphic form or 
possible sibling species). 

Callophrys (Mitoura) gryneus gryneus (Hubner, 1819), L-m-a-r.” 

C. (M.) gryneus auburniana (Harris, 1862), Bastrop Co. (Sandy Cr.). 

C. (M.) gryneus X sweadneri, Travis Co. (3 in hundreds of the parents). 


. C. (M.) sweadneri (Chermcck, 1944) subsp., Q-e-a-r.¥ 
. C. (UIncisalia) solatus (Cook & Watson, 1909), Q-n-a-r (with a var. with green 


scales ventrally).* 
C. (I.) henrici turneri (Clench, 1943), Llano, Travis Co’s. 


. *Atlides halesus estesi Clench, 1942, D-m-a-r. 

. Fixsenia ontario autolycus (Edwards, 1871), B-m-a-r. 

. Panthiades m-album m-album (Boisduval & LeConte, 1833), L-sw-u-r. 
. *Strymon melinus franki Field, 1938, C-e-a-r (with a dark winter f.).% 
. S. melinus melinus Hubner, 1818, N-w-u-r. 

. S. alea (Godman & Salvin, 1887), P-n-a-r (with a dark winter f.).'® 

. S. alea X columella (1 in 75 specimens of parent species). 

. S. columella istapa (Reakirt, 1866), R-n-s-p (14:x:71; 2:x:72; 7, 9, 10:xi:72). 
. Hemiargus ceraunus zachaeina (Butler & Druce, 1872), D-e-u-p. 

. Echinargus isola alce (Edwards, 1871), C-m-a-r (with dark winter f.). 

. Celastrina ladon ladon (Cramer, 1780), M-s-s-r (6:viii:72).17 

. Zizula cyna (Edwards, 1881), R-n-s-p (2:xi:71). 

. Brephidium exilis exilis (Boisduval, 1852), B-e-s-r (22:x:71; 2, 4:xi:71). 
. Everes comyntas comyntas (Godart, 1824), H-w-u-p. 

. E. texana Chermock, 1944, Q-n-a-r.18 

. Leptotes cassius striatus (Edwards, 1877), R-n-s-p (17:v:72). 


L. marinus (Reakirt, 1868), D-e-u-p. 


PAPILIONINAE (PAPILIONIDAE) 


62. 


63. 
64. 
65. 


Papilio polyxenes curvifasci Skinner, 1902, B-m-a-r. 

P. rudkini f. clarki Chermock & Chermock, 1937, Travis Co. (Turkey Bend).!® 
Pterourus (Pterourus) troilus troilus (Linné, 1758), Travis Co. (Bull Cr.). 

P. (Jasoniades) glaucus glaucus (Linné, 1764), L-sw-u-r. 

P. (J.) multicaudatus (Kirby, 1884), O-e-u-r (with large dark summer f.). 
Heraclides (Heraclides) cresphontes (Cramer, 1777), G-m-a-r. 

H. (H.) thoas autocles (Rothschild & Jordan, 1906), Travis Co. (L. Austin). 


66. *Battus philenor philenor (Linné, 1771), C-m-a-r. 
B. polydamas polydamas (Linné, 1758), Travis Co. (Austin). 
LIBYTHEIDAE 
67a. Libytheana bachmanni bachmanni (Kirtland, 1851), L-sw-u-p (with wet season 


67b. 
68. 


f. kirtlandi (Field, 1938)).?° 

*L. bachmanni larvata (Strecker, 1878), D-n-a-r (with wet season f. streckeri 
(Field, 1938)). 

L. carinenta mexicana Michener, 1943, R-n-s-p (3:v:73; 24:ix:79; 2:x:80; 
11:xi:80). 


NYMPHALINAE (NYMPHALIDAE) 


. *Euptoieta claudia claudia (Cramer, 1776), C-m-a-r (with winter dwarfs). 
. E. hegesia hoffmanni Comstock, 1944, R-n-s-p (16, 21:x:71). 
. Polygonia (Polygonia) interrogationis (Fabricius, 1869), K-m-a-r (with summer 


f. umbrosa (Lintner, 1869)). 


. P. (Grapta) comma (Harris, 1842), M-sw-s-r (with summer f. dryas (Edwards, 


1870)), (24, 26:ix:79). 


. Nymphalis (Euvanessa) antiopa lintnerii (Fitch, 1856), O-e-s-r (6:ix:76). 

. *Vanessa (Vanessa) atalanta rubria (Fruhstorfer, 1916), A-m-a-r. 

. *V. (Cynthia) cardui (Linné, 1758), A-m-a-p. 

. *V. (C.) virginiensis (Drury, 1773), A-m-a-r (with winter f. fulvia (Dodge, 1900)). 


VOLUME 36, NUMBER 1 le 


th 
78. 


79a 


79b. 
80. 


81. 
82. 


83. 
84. 


96. 


*Precis (Junonia) coenia coenia (Hubner, 1822), C-m-a-r (with dark wet season 
f., and winter f. rubrosuffusa Field, 1936). 

P. (J.) nigrosuffusa (Barnes & McDunnough, 1916), D-n-u-p.”! 

P. J.) genoveva genoveva (Stoll, 1782), R-n-u-p. 

P. (J.) genoveva zonalis (Felder & Felder, 1867), R-n-s-p (2:x:71; 2:xi:71; 
1-vii:72). 

Anartia jatrophae jatrophae (Johansson, 1763), R-nm-s-p (18:x:71). 

Siproeta (Victorina) stelenes biplagiata (Fruhstorfer, 1907), Travis Co. (Austin; 
R. Neck). 

Heliconius charitonius vazquezae Comstock & Brown, 1950, R-nm-a-p (with var. 
with orange scaled disc). 

Eueides isabellae zorcaon (Reakirt, 1866), Travis Co. (Austin, R. Neck). 

Dryas iulia moderata (Riley, 1926), R-nm-a-p (with dark wet season f.). 

Dione (Dione) moneta poeyi (Butler, 1873), Travis Co. (Bull Cr.; P. Horde). 
*D. (Agraulis) vanillae incarnata (Riley, 1926), B-m-a-r (with winter dwarfs). 
Mestra hypermnestra amymone (Ménétriés, 1857), R-m-a-p (with wet season 
dark f.). 

Biblis hyperia aganisa Boisduval, 1836, Travis Co. (Austin). 

Chlosyne (Chlosyne) janais (Drury, 1782), Travis Co. (Bull Cr.). 


. C. (C.) lacinia adjutrix Scudder, 1875, J-m-a-r (with dry season light f. cf. saun- 


dersi (Doubleday, 1848), and var. black, var. black and white). 


. C. (Charidryas) gorgone carlota (Reakirt, 1866), B-s-s-p (5:iv:78; 29:ix:79). 
. C. (C.) nycteis (Doubleday, 1848), M-sw-a-r (with wet season f. cf. drusius (Ed- 


wards, 1884)). 
Thessalia theona bollii (Edwards, 1877), Burnet Co. (Pangle). 


. Texola elada ulrica (Edwards, 1877), E-n-a-r (with wet season f. senrabii (Barnes, 


1900)). 


. *Phyciodes phaon (Edwards, 1864), L-m-a-r (with summer f. aestiva (Edwards, 


1878)). 


. *P. tharos distincta Bauer, 1975, O-e-a-r (with winter f. ventrally brown). 
. *P. vesta (Edwards, 1869), J-e-a-r (with winter f. boucardi Godman & Salvin, 


1878). 


. *Anthanassa texana texana (Edwards, 1863), J-m-a-r (with summer f. smerdis 


(Hewitson, 1864), and var. black & white). 


. Marpesia (Euglyphus) chiron (Fabricius, 1775), R-nm-s-p (29:ix:75). 

. Limenitis (Limenitis) bredowii eulalia (Doubleday, 1848), O-e-a-p.”” 

. L. (Basilarchia) archippus archippus (Cramer, 1776), B-s-a-p. 

. L. (B.) archippus watsoni (dos Passos, 1938), N-w-a-r (with var. orange, a Bates- 


ian mimic of Danaus eresimus v. orange).”° 
L. (B.) astyanax astyanax (Fabricius, 1775), L-sw-u-r. 
Dynamine dyonis Geyer, 1837, Travis Co. (Bull. Cr.). 


APATURINAE (NYMPHALIDAE) 


97. 
98. 
99. 
100. 
. A. louisa Stallings & Tumer, 1947, P-n-u-r. 
. *Anaea (Anaea) andria andria Scudder, 1875, L-m-a-r (with winter f. andriaesta 


103. 


Asterocampa celtis alicia (Edwards, 1868), N-w-a-r. 

A. antonia antonia (Edwards, 1877), B-m-a-r. 

A. leila cocles (Lintner, 1884), F-n-a-p. 

A. clyton texana (Skinner, 1911), B-m-a-r (with light dry season f.). 


Johnson & Comstock, 1941). 
*A. (A.) aidea (Guerin, 1844), R-nm-a-p (with winter f. morrisonii (Edwards, 
1883)). 


SATYRINAE (NYMPHALIDAE) 


104. 


Cercyonis pegala texana (Edwards, 1880), B-m-a-r (with var. dark). 
Cyllopsis gemma gemma (Hubner, 1808), Lee Co. (Patschke Bog). 
Neonympha (Hermeuptychia) hermes (Fabricius, 1775), Travis Co. 


12 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


105. N. (H.) sosybius (Fabricius, 1793), N-w-a-r.”* 

106. Megisto cymela cymela (Cramer, 1777), B-m-a-r (with var. smeared silver). 

107. M. rubricata rubricata (Edwards, 1871), B-m-a-r (with var. obsolescent eye- 
spots). 


DANAINAE (NYMPHALIDAE) 


108. Danaus (Danaus) plexippus plexippus (Linné, 1758), A-m-a-p (with var. dark 
apex forewing). 

109. *D. (Anosia) gilippus strigosus (Bates, 1864), C-m-a-r (with var. cf. berenice 
(Cramer, 1780), var. gilippina Hoffmann, 1940, and var. orange Mullerian mimic 
of D. eresimus var. orange). 

110. D. (A.) eresimus montezuma Talbot, 1943, R-n-a-p (with var. dark maculate, and 
var. orange). 


PIERINAE (PIE RIDAE) 


111. Anthocharis (Paramidea) midea annickae dos Passos & Klots, 1969, M-sw-a-r. 
112. Artogeia rapae rapae (Linné, 1758), A-m-a-p (exotic introduction). 
113. *Pontia protodice (Boisduval & LeConte, 1829), A-m-a-r (with winter f. vernalis 
(Edwards, 1864)). 
114. Ascia (Ascia) phileta phileta (Fabricius, 1775), R-nm-s-p (16:x:71).”° 
115. Appias (Glutophrissa) drusilla poeyi (Butler, 1872), R-nm-s-p (13, 18:x:71). 
116. *Colias (Colias) eurytheme eurytheme Boisduval, 1852, A-m-a-r (with summer 
f. amphidusa Boisduval, 1852, and female var. alba Strecker, 1878). 
C. (C.) philodice philodice Godart, 1819, Bastrop Co. (H. Duval Cln.). 
117. *Zerene cesonia cesonia (Stoll, 1790), A-m-a-r (with winter f. rosa (M Neill, 
1889), and var. stainkeae Field, 1936). 
Anteos maerula lacordairei (Boisduval, 1836), Travis Co. (N. Austin). 
118a. *Kricogonia lyside lyside (Godart, 1819), D-e-a-p (with winter f. unicolor God- 
man & Salvin, 1889).?° 
118b. K. lyside terissa (Lucas, 1852), R-nm-a-p (with f. lanice Lintner, 1885). 
118c. K. lyside fantasia Butler, 1871, R-n-u-p. 
119. Eurema (Eurema) mexicana (Boisduval, 1836), O-e-a-r (with winter f. rosa Whit- 
taker & Stallings, 1944). 
E. (E.) daira daira (Godart, 1819), Travis Co. (Austin). 
120. E. (Pyrisitia) proterpia (Fabricius, 1775), R-n-a-p (with winter f. gundlachia 
(Poey, 1851)). 
121. E. (P.) lisa Boisduval & LeConte, 1829, C-m-a-r (with summer f. immaculata 
Whittaker & Stallings, 1944, and female var. alba Strecker, 1878). 
122. E. (P.) nise nelphe (Felder, 1869), R-n-s-p (14:x:76; 11:xi:80). 
123a. *E. (Abaeis) nicippe nicippe (Cramer, 1780), C-m-a-r (with winter f. ventrally 
orange, and female var. pale orange). 
123b. E. (A.) nicippe flava (Strecker, 1878), N-w-s-r (19:x:71).?7 
124a. Phoebis (Callidryas) sennae eubule (Linné, 1767), L-sw-a-p (with female var. 
browni Field, 1936). 
124b. P. (C.) sennae sennae (Linné, 1758), N-w-a-p. 
124c. P. (C.) sennae marcellina (Cramer, 1777), R-nm-a-p (with female var. yamana 
Reakirt, 1863).78 
125. P. (C.) philea (Johansson, 1763), R-m-s-p (10:x:71; L. Gilbert). 
126. P. (Phoebis) agarithe maxima (Neumoegen, 1891), R-m-a-p (with female var. 
albarithe Brown, 1929). 
Aphrissa statira statira (Cramer, 1777), Travis Co. (Northwest Hills). 
127. *Nathalis iole Boisduval, 1836, A-m-a-r (with winter f. viridis Whittaker & Stall- 
ings, 1944). 


FOOTNOTES TO LIST OF BUTTERFLIES ON BARTON CREEK 


‘Hesperia tamenund Edwards has been treated as a subspecies of Papilio thraso Jung, 1792 which is a junior 
homonym. Papilio mithridates Fabricius, 1793 is the oldest name for the species (H. Ebert, 1969, J. Lepid. Soc. 23, 
Suppl. 3: 38). 


VOLUME 36, NUMBER 1 13 


2 Nisoniades llano Dodge, described from Llano Co., refers to the Central Texas populations which are subspe- 
cifically distinct from Thanaos invisus Butler & Druce, 1872 which ranges north to Tamaulipas and strays into the 
Rio Grande Valley. 

3 Species allied to Papilio oileus Linné, fall outside the genus Pyrgus as revised by B. C. S. Warren (1926, Trans. 
Entomol. Soc. London, 74: 152). By genitalic structure they fit best in Tuttia Warren, 1926. This genus was sunk 
in Muschampia Tutt, 1906 (e.g., C. G. Higgins & N. D. Riley, 1970, Field Guide to the Butterflies of Britain & 
Europe. Boston) for which F. Hemming (1967, Bull. British Museum Nat. Hist., Entomology, Suppl. 9: 300) has 
shown the correct name is Syrichtus Boisduval, 1834. 

4 Pyrgus albescens Plotz is distinguished genitalically from Syrichthus communis Grote, as are occasional inter- 
mediates interpreted as hybrids. Genitalic determination of males is made in the field using a 20x Seibert emoskop. 
A. W. Lindsey, E. L. Bell & R. C. Williams (1932, J. Sci. Lab., Denison Univ., 26: 46) are followed in recognizing 
subspecific status. 

> Specific distinctness of Papilio oileus Linné and Pyrgus philetas Edwards has been supported on structural, 
ecological and geographic grounds by J. M. Burns & R. O. Kendall (1969, Psyche, 76: 453). 

®H. A. Freeman (1969, J. Lepid. Soc. 23, Suppl. 1) reports different chromosome numbers (27) for Megathymus 
coloradensis (Riley, 1877) and (26) for M. yuccae (Boisduval & LeConte, 1833) and places M. y. reinthali Freeman, 
1963 with coloradensis rather than with yuccae. 

7 Pamphila licinus Edwards, 1871, described from Waco (Belfrage clr.) although recently treated subspecifically 
under Hesperia metea Scudder, 1863 by MacNeill (in Howe et al., 1975, Butterflies of North America. New York) 
is considered distinct following Lindsey, Bell & Williams (loc. cit.) until data is available from connecting localities. 

8 This subspecies occurs on southern extensions of tall grass prairie developed as glades on fluvial terraces on 
and west of the Balcones Fault Zone. The next subspecies occurs on Fayette prairie, a facies of the Southeastemm 
Coastal Plain prairies. The first is identified with Hesperia iowa Scudder (type locality: Denison, Iowa); the second 
with H. arogos Boisduval & LeConte (type locality: “North America,” probably Georgia), determination and sub- 
specific ranking following A. B. Klots (1951, Field Guide to the Butterflies. Boston) rather than MacNeill (loc. cit.). 

’ Subspecific recognition follows W. S. McAlpine (1971, J. Res. Lepid., 10: 28) rather than the lumping of J. A. 
Powell (in Howe et al., loc. cit.). The allolectotype, fig. 6 of F. M. Brown (1968, Trans. American Entomol. Soc., 94: 
130) is of the summer form. 

1 Here the synonymy of Powell (loc. cit.) is followed with added recognition that guadeloupe was based on the 
winter form more easily confused with the summer form of C. nemesis. C. rawsoni McAlpine, 1939 was based on 
the distinctive summer form of this species. Although convenient labels for ecologically important phenotypes, 
these names, when used for seasonal forms, are infraspecific and outside present zoological nomenclature. 

1 Specimens collected by Morgan Hebard, 23 miles east of Austin (Carnegie Museum Cln.) fall close to mejicanus 
and are the basis for this determination. One sight record (Travis Co., Austin, Northwest Hills, 13:x:68) resembled 
the Hebard specimens. 

2 W.T. M. Forbes (1960, Cornell Univ. Agric. Expt. Sta., Mem. 371: 133) is followed in recognition of subspecific 
distinctness of the northeastern entity associated with red cedar. Bastrop Co. material (associated with Juniperus 
virginiana var. crebra Fem. & Grisc.) was compared with series from Connecticut, New Jersey and North Carolina. 
This is distinct in color facies from the southern subspecies gryneus which is synonymous with smilacis Boisduval 
& LeConte, 1833. Texas material of this complex is routinely referred to castalis Edwards, 1871 (type locality: 
Waco) but the type seems indistinguishable from the summer form of southeastern gryneus. Our gryneus is asso- 
ciated both with Juniperus silicicola (Small) Bailey and with J. ashei Buchh. in lowland sites, perhaps favoring 
hybrid stands of these species. 

13 Our other species of C. (Mitoura) is associated with the upland phenotype of J. ashei which resembles J. 
monticola Martinez. Specimens were compared with the type series of C. sweadneri with which our winter brood 
is in close agreement. One specimen of our summer brood at Carnegie Museum (Austin, an old collection, possibly 
by C. T. Brues or J. F. McClendon) was placed under C. siva which it superficially resembles. K. Johnson (1978, 
J. Lepid. Soc., 32: 3-19) has not recognized three taxa in Central Texas. 

44 Incisalia henrici var. solatus Cook & Watson (type locality: Blanco Co., Texas) originally proposed as a “geo- 
graphic variety” and formally accorded subspecific status by W. Barnes & J. H. McDunnough (1917, Checklist of 
the Lepidoptera of Boreal America. Decatur, II].) is here broadly sympatric, and often syntopic and synchronic with 
C. henrici turneri (Clench, 1943). The former normally utilizes buds, flowers and early pods of Sophora secundiflora 
as larval food, but when these are destroyed by late frost, it shifts to Diospyros texanum; the latter utilizes flowers 
and fruit of Cercis canadensis as larval food, with populations in Llano and Bumet Co. using Lupinus texensis 
Hook. Specific distinction is inferred from these differences and the lack of clearly intermediate individuals. 

5 As well as obviously seasonal variation, a suite of apparently genetically determined phenotypes occurs in our 
area. The most frequent of these is identified as S. m. franki (type locality: Lawrence, Kansas), and the next most 
frequent as S. m. melinus (type locality: coastal Georgia). The former is associated with drier and prairie sites; the 
latter with streamside herbaceous stands, particularly with Hypericum sp. 

16H. K. Clench (1966, J. Lepid. Soc., 20: 65-66) is followed for treatment of our material. There is still not enough 
west Mexico topotypical material to resolve the need or not for recognition of Callicista laceyi Barnes & Mc- 
Dunnough, 1910 as eastern subspecies, queried by Clench. 

“ The correct name for this entity, formerly known as Argus pseudargiolus Boisduval & LeConte, 1833 has been 
clarified by H. K. Clench & L. D. Miller (1980, J. Lepid. Soc., 34: 103=119) who resurrect the same opinion of A. 
G. Butler (1900, Canadian Entom., 32: 91). 

‘8 This entity is not a subspecies of E. comyntas which also occurs this far and farther south. It is probably 
conspecific with E. herri (Grinnell, 1901) but not with E. amyntula (Boisduval, 1852) but this remains to be properly 
demonstrated. 

18 The correct name for this entity is uncertain. Edwards (1877, Trans. American Entomol. Soc., 6: 10) was probably 
correct in considering Arizona material conspecific with P. americus Kollar, 1850. P. coloro Wright, 1906 may refer 
to the California subspecies currently called P. rudkini J. A. Comstock, 1935. All material seen from east of the 
continental divide and north of latitude 30° is of f. clarki. P. americus stabilis Rothschild & Jordan, 1906, which 
more closely resembles P. coloro, ranges north from eastern Mexico to Bexar and Comal Co’s. in Texas. Brown’s 
(1942) f. pseudoamericus is an aberrant P. polyxenes asterius quite unrelated to the entity in question. 

2° These three Libytheana taxa have been frequently confused (e.g., Howe, 1975, loc. cit.: 258, pl. 47, f. 14, 15, 


14 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


presents L. bachmanni larvata as L. carinenta mexicana). Heitzman & Heitzman (1972, J. Res. Lepid., 10: 284) 
give good figures for determination of the North American mainland species. 

21 Thorne (1971, J. Res. Lepid., 9: 101) is followed in treatment of the P. coenia complex and all males were 
genitalically determined. Work by Clench, continued by Harvey suggests that P. evarete (Cramer, 1780) is an 
additional species distinct from our three. Infrequent intermediates between P. g. genoveva and P. g. zonalis suggest 
conspecificity; genoveva occurs south of, and zonalis east of our area. De Lesse (1952, Bull. Soc. Ent. France, 57: 
74) is followed in recognition of Junonia but only at the subgeneric level. 

22 By genitalia and venation L. bredowii eulalia is close to the type species of Limenitis, L. populi (Linné, 1758). 
As demonstrated by G. D. H. Carpenter & B. M. Hobby (1944, Trans. Royal Entomol. Soc. London, 94: 311-346) 
it is not an Adelpha, that genus being closer to Parthenos. Species related to L. astyanax (Fabricius, 1775) have 
been separated generically from L. bredowii (e.g., dos Passos, 1964, Lepid. Soc. Mem. 1) but the available name 
Basilarchia Scudder, 1872 is here used subgenerically. 

237. a. archippus is the plains and northeastern subspecies which ranges south to the Rio Grande in West Texas. 
L. a. watsoni is the Gulf Coast subspecies which ranges west to the Rio Grande, meeting archippus in Travis and 
Zapata Co’s. Here populations of one or the other wax and wane with little intergradation. 

24 Neonympha Hubner, 1818: 8 (type species: Papilio areolatus Abbot in Smith, 1797) is senior by page to 
Euptychia Hubner, 1818: 20 (type species: E. mollina Hubner, 1818). W. Forster (1964, Veroffentlichungen der 
Zoologischen Staatssammlung, Munchen, 8: 88-89) is followed in specific distinction of hermes and sosybius, 
although his genus Hermeuptychia seems no more than subgenerically distinct from Neonympha. 

25 G. Talbot (1928, Bull. Hill. Museum, 2: 195) pointed out that Papilio monuste Linné, with which our species 
has been identified, is actually Udaina cycnis Hewitson. Papilio phileta Fabricius, 1775 is the oldest available 
name. 

26 The entities lyside, terissa, and fantasia have geographically recognizable ranges that overlap only in northeast 
Mexico and Texas, and then only during migration. N. D. Riley (1972, J. Lepid. Soc., 26: 228) determined that 
Papilio castalia Fabricius, 1793 is a junior synonym of P. drusilla Cramer, 1777, and the entity that has been known 
as castalia (e.g., J. L. de la Torre y Callejas, 1958, Univ. de Oriente (Cuba), Dept. Ext. y Relac. Cult., 42: 24) should 
be referred to the appropriate senior available name, Gonepteryx terissa Lucas, 1852. 

27 Holland (1932, The Butterfly Book. New York) figures a male of this entity (pl. 37, f. 5). L. Harris (1972, 
Butterflies of Georgia. Norman, Oklahoma) reports three collections from Georgia, including three males and two 
females. Originally described as a species, in recent usage (e.g., C. P. Kimball, 1965, Lepidoptera of Florida. 
Gainesville) this name has been erroneously applied to the light var. of the orange female from which the true flava 
may be distinguished by the cold yellow tone resembling E. boisduvaliana Felder & Felder, 1865 with which it 
may be confused on the wing. 

28 Some years eubule, sennae, and marcellina are all absent. In other years a combination of one to rarely three 
are present. The low frequency of intermediates between eubule and the other two denies a clinal relationship and 
may ultimately prove specific distinction. 


Faunal history. Micropaleoentomological studies in progress in- 
dicate that in this region, pluvial and interpluvial climates were out 
of phase with glacial and interglacial climates of the north. River ter- 
race deposits (largely gravel) appear to represent interpluvial time. 
They were deposited when sea level stood higher than today and 
river gradients were adjusted upward. Accumulation took place dur- 
ing infrequent severe storms in a period of semiarid climate during 
later interglacial and early glacial times. Interpluvial times appear to 
have occupied much more of the Pleistocene Epoch than did pluvial 
times. The former were times of plains and desert communities in 
our area and of isolation of relict pockets of woodland biota. Pluvial 
times were brief episodes of dispersal between relict woodlands and 
influx of Sierra Madre montane and eastern deciduous forest elements 
during cool pluvial time, and tropical forest elements during warm 
pluvial time. Periodic colonists are of course in a state of flux, depen- 
dent upon minor climatic fluctuations of the recent past. Our resi- 
dent species may be analyzed in terms of a tentative schedule of late 
Pleistocene episodes based on data published for Hershop, North 
Soefje, and Rutledge Bogs in Gonzales Co., Boriack and Patschke 
Bogs in Lee Co., South Gause and North Gause Bogs in Milam Co., 
and Franklin Marsh in Robertson Co. (Durden, 1979). 


VOLUME 36, NUMBER 1 15 


For the last 4000 years our regional fauna has probably been much 
the same as present under moderate and dry climate with reduction 
of periodic colonists from the south during the “Little Ice Age”. From 
6000 to 4000 years ago the climate was moderate and not quite so 
dry. A few periodic colonists may have assumed resident status. From 
8000 to 6000 years ago the climate was moderate and nearly as dry 
as present, with presumably almost the same fauna. From 9200 to 
8000 years ago the climate was moderate and moist, with probable 
resident status for a few of our southern and eastern periodics. From 
9700 to 9200 years ago the climate appears to have been hot and dry. 
The Austin regional fauna at this time probably resembled that of the 
lower Rio Grande Valley with 270 or more species of butterflies. 
Our relict populations of species with Tamaulipan (Durden, 1974) 
affinities probably date from this time. These include Systasea pul- 
verulenta, Ancyloxipha arene, Apodemia mormo mejicana, Strymon 
alea, Thessalia theona bollii, Texola elada ulrica, Phyciodes vesta, 
Asterocampa louisa, and Neonympha hermes. The Travis Co. (Turkey 
Bend) population of Papilio rudkini f. “clarki” probably arrived from 
the Chihuahuan and Sonoran Deserts at the same time. From 
11,500 to 9700 years ago the climate was warm and arid, and 
some of the previously mentioned species may have established 
residency this early. From 12,800 to 11,500 years ago the climate 
was cooler than the present and dry. Our resident species of 
Gulf Coastal prairie and savanna distribution probably date from 
this episode. These include Thorybes confusis, T. bathyllus, Eryn- 
nis zarucco, Hesperia meskei, Atrytone arogos arogos, Ambly- 
scirtes alternata and Cyllopsis gemma gemma. Species of Great 
Plains distribution probably date largely from this time also. These 
include Pholisora catullus, Copaeodes aurantiaca, Hesperia viridis, 
H. attalus, Atrytone arogos iowa, A. delaware lagus, Amblyscirtes 
nysa, A. eos, Phaeostrymon alcestis, Strymon melinus franki, and 
Brephidium exilis exilis. From 14,000 to 12,800 years ago the climate 
was warm and very wet. We appear to have no tropical species sur- 
viving here from this episode, but some of the Gulf Coast swamp- 
woodland species may go back this far. These include Syrichtus 
oileus, Strymon melinus melinus, Limenitis archippus watsoni, As- 
terocampa celtis alicia, and Eurema nicippe flava. At this time dwarf 
palmetto reached Barton Creek where it is now disjunct some 100 km 
from the nearest Coastal Plain stand down river. From 15,000 to 
14,000 years ago the climate was moderate and wet. From 16,000 to 
15,000 years ago and possibly as early as 22,000 years ago the climate 
was cool and moist. This appears to be the episode from which date 
both our relicts of Virginian distribution and of Sierra Madre montane 


16 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


distribution. The former include Thorybes pylades, Erynnis meridi- 
anus, E. juvenalis juvenalis, E. martialis, E. baptisiae, Ancyloxipha 
numitor, Polites themistocles, Poanes viator, Feniseca tarquinius, 
Satyrium calanus falacer, Callophrys gryneus auburniana, C. henrici 
turneri, Panthiades m-album, Celastrina ladon, Pterourus troilus, P. 
glaucus, Polygonia comma, Chlosyne nycteis, Limenitis astyanax, 
Neonympha sosybius and Anthocharis midea annickae. The latter 
include Epargyreus clarus subsp., Achalarus coyote, Cogia outis, 
Gesta invisus llano, Hesperia woodgatei subsp., Amblyscirtes aenus, 
A. erna, A. celia, Calephelis guadeloupe, C. sinaloensis nuevoleon, 
Callophrys sweadneri subsp., C. solatus, Pterourus multicaudatus, 
Nymphalis antiopa lintneri, Siproeta stelenes biplagiata, Phyciodes 
tharos distincta and Eurema mexicana. However, many of these ap- 
pear to have persisted in our area over several interglacials, as have 
such species of similar affinities as Megathymus coloradensis ken- 
dalli, M. c. reinthali, Hesperia licinus, Calephelis nemesis australis, 
C. perditalis and Everes texana. Betore 22,000 years ago there appear 
to have been long episodes of climate much cooler than present and 
rather arid. Some of our species of Great Plains affinities may go back 
this far, but the only one that certainly dates from this time is the 
Blanco Co. (Round Mountain) population of Erynnis burgessi. Our 
fauna at that early and mid-glacial time must have been impoverished, 
resembling that of the Davis or Guadalupe Mountains with their mon- 
tane endemic elements represented by our own canyon relicts. 

The butterfly fauna of the Austin area is the result of opportunistic 
colonization under each changed climate, with the persistence of rel- 
ict populations in reduced habitats through subsequent unfavorable 
climatic episodes. Our endemic taxa, most of which are shared with 
the mountains of northern Coahuila, Mexico, are the few lineages that 
have persisted through much of the Pleistocene and probably longer. 
The community to which they belong includes relict plants such as 
Berberis swaseyi Buckl. which find their roots in the Oligocene biota — 
of Florissant, Colorado. 

Potentially threatened or endangered taxa. None of the 
species that occur in the study area appear to be threatened by normal 
collecting for scientific or recreational purposes. The greatest hazard 
to all species of exacting habitat requirements is destruction of hab- 
itat. One of the six known stations for Calephelis sinaloensis nuevo- 
leon in the United States is now an apartment complex and parking 
lot with no butterflies. This species with only two known additional 
stations in Mexico should be watched closely. The study site on Bar- 
ton Creek is a protected station that should be preserved by non-« 
development of the brush and chaparral where Eupatorium hava- 


VOLUME 36, NUMBER Il IG 


nense, the presumed larval food plant, grows. Bernardia myricaefolia, 
the larval foodplant of Strymon alea, is equally vulnerable to habitat 
destruction. The grassland species, Hesperia woodgatei and Everes 
texana, are susceptible to loss of foodplant through grazing. One col- 
ony of the latter in Travis Co. (Bull Creek) is gone through overgraz- 
ing by horses. 


ACKNOWLEDGMENTS 


This project was stimulated by work with Harry Clench at Powdermill Nature Re- 
serve of Carnegie Museum from 1965 to 1968. Harry was responsible for addition of 
a species to this list during a visit in 1972 (Artogeia rapae!). Lawrence E. Gilbert, 
Raymond Neck, Roy O. Kendall, William W. McGuire, Pete Hord, and Donald J. Har- 
vey have provided information and field company on occasion. Determinations, habi- 
tats and ranges were discussed with all these colleagues but the data presented here 
is the sole responsibility of the author. Systematic descriptions are in preparation for 
a few new taxa indicated but not named here. A phenography or calendar of adult flight 
times will be published shortly at Texas Memorial Museum. 


LITERATURE CITED 


CORRELL, D. S. & M. C. JOHNSTON. 1970. Manual of the Vascular Plants of Texas. 
Texas Research Foundation, Renner. 1881 pp. 
DURDEN, C. J. 1969. Ecological aspects of the species concept. Armadillo Papers 1. 14 


Pp. 


1974. Biomerization: An Ecologic Theory of Provincial Differentiation, in C. 

A. Ross, ed., Paleogeographic Provinces and Provinciality. Soc. Econ. Paleontol. 

& Mineral., Spec. Publ. 21: 18-53. 

1979. Moss, sedge and forest bogs of Texas. Texas Natural Areas Survey (un- 
published report). 36 pp., map. 

HOLDRIDGE, L. R., W. C. GRENKE, W. H. HATHEWAY, T. LIANG & J. A. Tost. 1971. 
Forest Environments in Tropical Life Zones. Pergamon, Oxford. 780 pp. 

NECK, R. W. 1978. Climatic regimes resulting in unusual occurrences of Rhopalocera 
in Central Texas in 1968. J. Lepid. Soc., 32: 111-115. 

Roppa, P. U., L. E. GARNER & G. L. DAWE. 1970. Austin West, Travis County, Texas. 
Univ. Texas, Austin, Bur. Econ. Geology, Geol. Quadr. Map 38, + 11 pp. 

U.S. DEPT. COMMERCE, NATIONAL OCEANIC AND ATMOSPHERIC ADMINISTRATION, 
ENVIRONMENTAL DATA SERVICE. 1970. Local Climatological Data, Austin, Texas. 
4 pp. 

WERCHAN, L. E., A. C. LOWTHER & R. N. RAMSEY. 1974. Soil Survey of Travis County, 
Texas. U.S. Dept. Agr., Soil Cons. Serv. & Texas Agr. Exper. Sta. 123 pp., 84 maps. 


EPILOGUE. Since May 1980 Thessalia theona has been taken at the study site and 
Pterourus palamedes (Drury, 1770) has been seen in Austin, raising the known local 
fauna to 128 species and the regional fauna to 173 species. Development adjacent 
to the study site has caused silting of the waterholes, massive bacterial contamination 
of the springs, and loss of cliff habitat replaced by condominium projects. 


Journal of the Lepidopterists’ Society 
36(1), 1982, 18-30 


THE LARVA AND STATUS OF CATOCALA PRETIOSA 
(NOCTUIDAE), WITH DESIGNATION OF 
A LECTOTYPE 


DALE F. SCHWEITZER 
Peabody Museum, P.O. Box 6666, Yale University, New Haven, Connecticut 06511 


ABSTRACT. A lectotype is designated and illustrated for Catocala pretiosa Lint- 
ner (1876) and the larva is described, illustrated, and compared with that of related 
species. The status of this taxon as distinct from C. mira Grote and C. crataegi Saunders 
is confirmed. However, its relationship to C. texarkana Brower (1976) is unresolved. 
The known present and former range of C. pretiosa is documented. Larvae were reared 
on Prunus maritima, a likely foodplant in New Jersey. However, no larvae could be 
located in the field. 


Catocala pretiosa Lintner (1876) was described from “three ex- 
amples ... captured by me at sugar, at Schenectady, N.Y., last year.” 
Dates given are 8 and 10 July (males), 16 July (female). The New 
York State Museum (NYSM) has a male in its type collection (as of 
December 1979) with three labels: 1) “Schenectady, N.Y. July 10, 
1874. Lintner Coll.”’; 2) “pretiosa Lintner 3966 3”; 3) “J. A. Lintner 
collection.” An enlarged photograph of the labels shows that the first 
printed label originally read 1875 but that a 4 was written in ink over 
the 5. The U.S. National Museum also has two specimens with the 
same locality and collector dated 15 July 1874 (female) and 8 July 
1875 (male). These labels are printed in the same style as that on the 
“type” and the 4 on the female’s label is similarly written over a 
printed 5. 

It is quite possible that the manuscript was written in 1875 and that 
“last year’ referred to 1874. It is also possible that Lintner was mis- 
taken about the years. At any rate, there can be virtually no doubt that 
the above NYSM specimen was among Lintner’s three specimens. 
Unfortunately, no mention of any types is made in the original de- 
scription. McCabe & Johnson (1980) listed this specimen as a syntype. 
I hereby designate this specimen, illustrated in Fig. 3, as Lectotype 
for Catocala pretiosa Lintner and will have an appropriate label af- 
fixed. 

Catocala pretiosa Lintner has remained a little known taxon since 
its description. Some authors (e.g., Barnes & McDunnough, 1918) 
have treated it as a form of C. crataegi Saunders. Forbes (1954) treated 
it as a species. Sargent (1976) reached no definite conclusion regard- 
ing its status. One of the major problems faced by taxonomists has 
been a shortage of specimens, especially recent ones. Quite a few old 
specimens lack data. 


VOLUME 36, NUMBER | 19 


Fic. 1. Mature larva (dorsal) of Catocala pretiosa, ex ovis Cape May, New Jersey, 
reared on Prunus maritima May-June 1978 by D. F. Schweitzer (2.54x). Larva at 
YPM. 


Based on material that I have seen and the literature (e.g., Sargent, 
1976; Brower, 1974), this species seems to have formerly been widely 
distributed in the northeastern United States, though it may have 
been partial to coastal areas (e.g., Stonington, Connecticut and Kittery, 
Maine) and sandy areas [e.g., the Merrimack River Valley in New 
Hampshire and the Albany-Schenectady (Centre), New York “Pine 
Bush’]. Bailey's account (1877) suggests it was locally common at 
Centre, New York. Despite this, nothing has been published regard- 
ing its life history or foodplant; so, the following description is pre- 
sented. About 13 late instar larvae reared from two females, both from 
Cape May, New Jersey taken in 1977 and 1978, were examined. 


Description of Early Life Stages 


Penultimate and ultimate instar larvae. As illustrated (Figs. 1, 2), dorsum pale gray 
and bold mid-dorsal stripe. Stripe, brown with thin darker, almost black, edges which 
do not include tubercles I or II. Facial stripe, black. Other dorsal markings, nearly 
untraceable. “Saddle” with faint light brown shading extending ventrad to tops of third 
and fourth prolegs. Venter whitish with very dark brown patches as shown. Dorsal 
horn present, brown. Dorsal ground color close to that in the black and white photo- 
graph in Fig. 1. 

Earlier instars. Ground color darker, with usual pattern more visible, middorsal 
stripe not darkened but with fragments of darker edging. 

Egg. Typical for the group, quite flat and rather circular. 

First, last and two intermediate instar larvae; eggs and pupal shells are preserved at 
Peabody Museum. Reared moths are in that collection and the author's. 


20 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Fic. 2. Same larva as Fig. 1 clinging to edge of glass showing ventral markings. 


Foodplant 


The larvae were reared on Prunus maritima, but the natural food- 
plant has not been established. Crataegus sp. (Pcrus-galli) was also 
accepted by hatchlings. Prunus maritima is abundant at the Cape 
May locality and is present at sites of recent captures of C. pretiosa 
at Atsion and Batsto, Burlington Co., New Jersey by myself and John 
Nordin. It does not occur at the immediate locality near Eldora, Cape 
May Co., New Jersey where a number of recent captures were made 
by Joseph Muller. P. maritima could not have been the host of the 
former inland C. pretiosa populations in New Hampshire and New 
York. Extensive searching and beating (including nocturnally) of P. 
maritima thickets on 7 and 20 May 1979 and 1 June 1980 at Batsto 
and Atsion failed to produce C. pretiosa larvae, although last and 
earlier instar Catocala ultronia Hubner larvae were found on both 
1979 dates. The few known Crataegus uniflora plants at Batsto were 
also checked in 1979. Prunus serotina is the only rosaceous tree or 
shrub known to be present at all New Jersey sites where C. pretiosa 
has been taken. Crataegus is scarce to absent at all such sites and in 
the New Jersey Pine Barrens region generally. Amelanchier and Py- 
rus (Aronia) spp. are frequent in this region. 


Taxonomic Status 


The notes on the larva of C. crataegi given by Forbes (1954) and 
the larval illustrations given by Barnes & McDunnough (1918) agree 


VOLUME 36, NUMBER 1 21 


Fic. 3. Lectotype Catocala pretiosa Lintner “Schenectady, N.Y. July 10, 1874. Lint- 
ner Coll.” “pretiosa Lintner 3966 3” “J. A. Lintner Collection” (New York State Mu- 
seum). 


well with each other and with Saunders’ (1876) original description. 
C. pretiosa differs in several aspects. C. pretiosa has a pale gray 
ground color; C. crataegi is decidedly brown. Catocala crataegi has 
a very different head capsule, featuring extensive brown mottling and 
a darkened face. C. pretiosa also lacks red or orange in the face which 
Saunders (1876) reports for C. crataegi. C. pretiosa also lacks orange 
on the tubercles and has a prominent brown line (not dots) on segment 
11 (not 12 as Saunders states). The horn is not red in C. pretiosa, and 
there are no green or blue hues ventrally, and the spiracles are dark. 
C. crataegi apparently does not have a form with a dark dorsal stripe. 

The alcohol preserved larva described as C. crataegi by Crumb 
(1956) appears to differ considerably from the above larvae and may 
be C. mira Grote. C. pretiosa differs from it on at least the following 
points: The dark granules are not more prominent on the thorax than 
on the abdomen; the middorsal stripe is dark brown, not pale, and its 
black border is quite prominent. The black U-mark on the face is not 
broken above seta A®. The adfrontal markings are very fine. 

The larva of C. mira reportedly has a long twisted horn (Barnes & 


22 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


McDunnough, 1918). I have collected such a larva in Connecticut (on 
Pyrus Xpurpurea); but several dozen known C. mira which I have 
seen from Florida had smaller untwisted horns as does Crumb’s 
(1956) larva. 

The larva of C. pretiosa differs from Crumb’s (1956) C. mira larva 
from Iowa on at least the following points: The middorsal stripe is 
dark; tubercles are not orange; spiracles are dark brown, not white; 
the head has little pattern aside from the U-mark, and no brown or 
protuberances. It differs from six C. mira larvae from Liberty Co., 
Florida (leg. H. D. Baggett & D. F. Schweitzer in YPM) in several 
details as well, including: the lack of protuberances on the face, which 
is also much less brown and paler on C. pretiosa; the lack of weakly 
defined transverse bands of brown flecks dorsally on the thoracic and 
first two abdominal segments; the less prominent brown on the saddle 
mark behind the horn; and the lack of orange on the tubercles. 

H. D. Baggett and colleagues have reared a number of C. texarkana 
Brower (most det. A. E. Brower) from larvae collected on Crataegus 
at Torreya State Park, Liberty Co., Florida from 1978 to 1980 (Fig. 9). 
I have examined photographs of larvae believed to be C. texarkana 
and they are similar to C. pretiosa. C. texarkana occasionally lacks 
the dark dorsal stripe and C. mira occasionally has such a stripe (H. 
D. Baggett, pers. comm. 1980, 1981). C. texarkana and C. mira feed 
together on Crataegus in April at Torreya State Park. C. mira, at least, 
also occurs there on wild plum (Prunus angustifolia) (reared by author, 
at YPM). 

I have reared three broods of Catocala blandula larvae from Con- 
necticut and one from Lebanon, New Jersey, and these were similarly 
dimorphic. These larvae differed from C. pretiosa in their lack of an 
abdominal horn and in having somewhat more dark mottling. C. pre- 
tiosa thus may prove to have an unstriped form. 

Catocala pretiosa adults differ from C. crataegi on a number of 
characters. C. pretiosa has a paler median area of the forewing, with 
the reniform much more conspicuously ringed with bright white. 
There is little or no darkening of the forewing inner margin of C. 
pretiosa, and little or no brown beyond the postmedian line on the 
forewing of C. crataegi. C. pretiosa has a deeper orange hindwing 
than C. crataegi. The pale median area can be used to separate both 
of these species from C. mira. The hindwing color of C. mira is like 
that of C. pretiosa. For further descriptions and color figures of this 
group see Forbes (1954), Sargent (1976) and Barnes & McDunnough 
(1918). 

Theodore Sargent has suggested to me that the present New Jersey 


VOLUME 36, NUMBER 1 2a 


Fic. 4. Catocala pretiosa 2, Schenectady, N.Y., 17 July 1877, J. A. Lintner coll. 
(NYSM). 


population (Figs. 5, 6) might not be the same as the original C. pre- 
tiosa of Lintner. Indeed my Batsto, New Jersey specimen that Sargent 
illustrated (1976, p. 67, B) is not typical of C. pretiosa. It differs most 
obviously in having the dark basal area on the forewing extending to 
the inner margin. However, this specimen appears to be atypical of 
the New Jersey population now that adequate material has been ex- 
amined. Most of the recent southern New Jersey specimens appear 
to me to match Lintner’s description, the lectotype and old New En- 
gland material quite closely. 

The forewing black area appears to be paler and browner on five 
of the seven old specimens I have at hand than on recent New Jersey 
ones (14, 9 reared, at hand). However, other old Catocala I have 
examined are frequently similarly discolored because of fading. 

Based on the above comparisons of larvae and the consistent dif- 
ferences in adult wing characters, I conclude that Catocala pretiosa 
is not conspecific with C. mira or C. crataegi. Catocala pretiosa is 
thus restored to its original status as a full species. 

Separation of Catocala texarkana from C. pretiosa is extremely dif- 
ficult. At present, some specimens cannot be determined with cer- 
tainty. The original description of C. texarkana does not contain ex- 


24 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


FIGs. 5, 6. Catocala pretiosa 3, @; siblings to larva in Figs. 1 & 2, eclosed 21, 27 
June 1978. (In author’s collection.) 


VOLUME 36, NUMBER | 25 


Fic. 7. Catocala ?texarkana 3; Chapel Hill, North Carolina, 12 June 1974, leg. 
Chas. C. Horton (in T. D. Sargent collection). 


plicit comparisons with related taxa, and I can find no constant 
differences between these two taxa. However, several characters will 
work for most specimens. 

Many C. texarkana var. “bridwelli” have the inner margin con- 
spicuously darkened in the median area as noted by Brower; almost 
all other specimens have some vague darkening of this region. C. 
pretiosa typically has only a few scattered dark scales, and no north- 
em specimens have this conspicuous dark median shade. Sargent 
(1976, p. 67, B, C) illustrates specimens showing both extremes for 
northern C. pretiosa. 

Typical C. texarkana has almost no black in the basal part of the 
forewing. A large C. pretiosa 2° from Sherborn, Massachusetts is es- 
sentially a perfect match for three such topotypical specimens of C. 
texarkana (YPM, Bryant Mather colls.). Most other C. pretiosa have 
extensive basal black or at least a mixture of brown and black scales 
(but see Fig. 4). C. texarkana form “‘bridwelli’” also has extensive 
basal black. 

The basal black of C. pretiosa stops abruptly at the anal vein as 
noted by Forbes (1954) but may continue (when present) to the inner 
margin in C. texarkana. The lone exception among C. pretiosa seen 
is my Batsto, Jersey ¢ illustrated by Sargent (1976, p. 67, B). 


26 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Fic. 8. Catocala P?texarkana 2°; Fontana Dam, Graham Co. North Carolina, 
1200-1800 feet, 8 July 1972, leg. D. F. Schweitzer (in author's collection). 


Most C. pretiosa have less darkening beyond the reniform than is 
present on C. texarkana. 

There apparently is some geographical variation in C. texarkana. 
Three Florida specimens and photographs of three others before me 
are all variety “bridwelli,’ and all have a prominently dark inner 
margin. Several other Florida specimens shown to me by H. D. Bag- 
gett were also “bridwelli.” This is apparently a minority form at the 
type locality (Brower, 1976). 

I have at hand a 6 from Chapel Hill (cited as C. pretiosa by Sargent, 
1976) and a pair from Fontana Dam, North Carolina. Two of these are 
illustrated (Figs. 7, 8). The Fontana 3 could pass for a rubbed C. 
pretiosa but has some of the median dark scaling at the inner margin. 
The female collected with it (Fig. 8) has a strongly darkened inner 
margin and the brightest white median area I have seen on either 
taxon. These three have less subapical brown shading in the post 
median gray than on any C. texarkana (three photographs, seven spec- 
imens) before me now—a trait shared by most C. pretiosa. Two of 
these also have less darkening beyond the reniform than is typical of 
either species, but some C. pretiosa agree closely. I see no point in 
placing allegedly certain names on them now. 

The specimens listed below from Virginia, Tennessee and Ohio all 


VOLUME 36, NUMBER 1 Ate 


Fic. 9. Catocala texarkana, dwarfed 6; Torreya State Park, Liberty Co. Florida, ex 
larva on Crataegus, eclosed 27 April 1978, leg. H. D. Baggett (in author's collection). 


have slightly more dark scales in the median white at the inner margin 
than more northeastern specimens. However, none has a solid dark 
inner marginal shade. These approach my Fontana ¢ (above). I can 
see no reason not to regard these specimens as C. pretiosa, despite 
this trivial difference. Texas, Florida, and North Carolina populations 
apparently are all composed largely of specimens that deviate more 
noticeably from northern C. pretiosa. 

More material from the southern Appalachians and other poorly 
collected southern regions is needed. Until such specimens are avail- 
able, the prudent course seems to be to treat the names in this group 
as proposed by their authors. Specimens from the Northeast can con- 
fidently be placed as C. pretiosa. Those from Ohio, Virginia, and 
eastern Tennessee appear almost identical. Populations in the Gulf 
Coast States appear to be C. texarkana. Specimens from North Car- 
olina are, at present, unplaceable. 


Distribution 


There is little chance that Catocala pretiosa occurs in upstate New 
York or New England at present. There have been no captures in 
those areas for at least 40, and probably 80, years despite fairly intense 
collecting. It has not been taken recently at Albany despite substantial 


28 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Fic. 10. Catocala:texarkana 2°; Torreya State Park, Liberty Co. Florida, 28 May 
1978, leg. D. D. Baggett (YPM). 


collecting by myself, T. L. McCabe, John Cryan and others. It also 
seems very likely that the species was formerly absent in southem 
New Jersey where it is now widespread and not rare. Lakehurst, New 
Jersey is one of the most intensively collected places in North Amer- 
ica and has been studied from before 1900 into the early 1970's, and 
C. pretiosa has not been taken there. The area is very similar to Atsion 
and Batsto, and Prunus maritima is frequent. Smith (1899, 1910) gives 
no southern New Jersey records for C. pretiosa. I have seen none in 
collections taken prior to 1968. Smith (1899) records C. crataegi from 
New Jersey near New York City, and states “The variety pretiosa 
Lint., has also been taken in the state.” 

The first record in southern New Jersey appears to be Joseph Mul- 
ler’s capture of three specimens at Cape May on 28 June 1968; sub- 
sequent captures have been from 1972 to 1981 at Atsion, Batsto, Elmer 
and Eldora by John Nordin, myself, the Rutgers University staff and 
Muller. The date range is 23 June to 16 July. I visited the exact Cape 
May locality in 1977 and 1978 and collected the species on each of 
three nights. The moths have been taken mostly at sugar baits. 

I have not attempted to catalog all records of C. pretiosa. However, 
during a visit to the U.S. National Museum (USNM) in July 1980, I 
recorded all specimens that seemed to be this species. These records, 


VOLUME 36, NUMBER 1 29 


plus the others given below, probably give a reasonable indication of 
the range of this species in the past. All recent records known to me 
are discussed above. All specimens recorded below were examined 
by me in 1979, 1980, or 1981 except as noted. 


NEW HAMPSHIRE: Milford, 10 July 1877, ex coll. C. P. Whitney, det. Lintner as 
typical pretiosa (not seen by me, record courtesy of Richard E. Gray, Montshire Mu- 
seum, Hanover, N.H.); Manchester (figured by Sargent, 1976, Pl. 8, Fig. 4; moth now 
at MCZ, Harvard U.). NEW YORK: Schenectady, July 10, 1874, Lintner coll., herein 
designated Lectotype (New York State Museum); Schenectady, July 15, 1974 and July 
8, 1875 (USNM); July 17, 1877 (NYSM) (all four ex Lintner coll.); Albany, July 4, 1877 
S. C. Waterman collector, ex Wm. W. Hill coll. (NYSM); Centre, July 1877, ex Oberthur 
coll. (USNM); Centre July 5, 1877 ex Oberthur & Barnes colls. (USNM); “Lint. N.Y.” 
(certainly Albany-Schenectady region leg. Joseph Lintner) (USNM); “coll. J. Angus, 
West Farms, New York City” (which may be merely an address label) (2 at Rutgers 
University and one, USNM); Duchess Co., 14 and 16 August (2, E. L. Quinter coll.); 
no specific locality ex Barnes coll. (USNM). MASSACHUSETTS: Sherborn, July ex E. 
J. Smith coll. (Yale Peabody Museum = YPM). CONNECTICUT: Stonington, 29 June 
1898 ex H. P. Wilhelm coll. (YPM, illustrated by Sargent, 1976, p. 67); no specific 
locality ex Barnes coll. (USNM); Ely coll. (no data, not definitely Connecticut, but very 
likely from East River, YPM); two with no data but believed to be from Connecticut, 
before 1940 ex John Reichert coll. (YPM). NEW JERSEY: no other data, “Col. B. 
Neumogen,’ ex Brooklyn Museum coll. (USNM); no specific locality July 83 ex J. B. 
Smith coll. (USNM); also Atsion, Batsto, Eldora, Elmer and Cape May, 1968-1980 (see 
above). PENNSYLVANIA: no specific locality, 10-6-96, ex Oberthur coll. (USNM). 
MARYLAND: no specific localities ex E. A. Smyth, Wm. Schaus & Edw. T. Owen colls. 
(USNM). OHIO: Columbus, W. N. Tallant coll., second label Edw. T. Owen coll. (3, 
(USNM); no specific locality, ex Wm. Schaus coll. (USNM). VIRGINIA: Montgomery 
Co., 1947, E. A. Smyth coll. (USNM). TENNESSEE: Norris Park, 27 May 1938 (2, J. 
W. Cadbury coll.); no locality, “Teneese” ex Dodge coll. (YPM); no specific localities 
5. 22, 27, 27, 28, all ex Bares coll. (USNM). 


Except for the southern New Jersey specimens, the records all ap- 
pear to represent very old specimens; the 1947 Virginia record is 
doubtless the most recent. Catocala pretiosa quite clearly was more 
frequent, at least northward, in the past than it is today. The Virginia 
specimen is probably the most northern capture between about 1920 
and 1968. 

Specimens seen from Tennessee, Virginia and Ohio differ from 
most of the others in having slightly more dark shading along the 
forewing inner margin, and some are somewhat browner (faded?) and 
larger. These specimens might be Catocala texarkana, but as noted 
above, I tentatively regard them as C. pretiosa. All specimens from 
Texas to Florida (see Figs. 8, 10) are presumably C. texarkana. 

Northern specimens should be easily identifiable, and at present 
C. pretiosa does not appear to be sympatric with either C. mira or C. 
crataegi. In New Jersey, C. pretiosa seems to be confined to the 
southern, coastal plain counties. I have one C. mira from this region 
(New Lisbon, 1972). Otherwise, C. mira and C. crataegi seem limited 
to the hilly northern counties. C. mira ranges from at least Massachu- 


30 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


setts to Florida and west through much of the Midwest. C. crataegi 
is decidedly northern, reaching southward to Hamden, Connecticut; 
Lebanon, Hunterdon Co., New Jersey (J. Muller); Schuylkill Co., 
Pennsylvania (E. L. Quinter); Chicago, Illinois (Crumb, 1956) and 
Louisiana, Missouri (USNM). I have seen the specimens cited by 
Kimball (1965) from Florida (USNM, AMNH) and these are similar 
to Baggett’s specimens of C. texarkana “bridwelli.” I have not seen 
C. crataegi from the southern Appalachians. 

If C. texarkana is in fact a good species and if the recent North 
Carolina specimens are C. texarkana, then southern New Jersey may 
be the last stronghold for C. pretiosa. However, this population seems 
to be of very recent origin, and perhaps other populations are still 
extant. This species should be looked for in the hard pine areas of 
eastern Maryland and southern Delaware. The prospects for long term 
survival of the species in southern New Jersey appear good if the 
foodplant is something common like Prunus maritima, which grows 
on coastal dunes and on disturbed sites and edges of woods in the 
Pine Barrens. 


ACKNOWLEDGMENTS 


I wish to thank H. D. Baggett for specimens and information concerning C. mira and 
C. texarkana in Florida and for taking me to the Torreya locality. I thank Charles L. 
Remington, Theodore D. Sargent and an anonymous reviewer for their suggestions for 
improving this paper. Timothy McCabe generously provided the photographs for Figs. 
3 and 4. All other photographs were taken by William Sacco, Peabody Museum, Yale 
University. 


LITERATURE CITED 


BAILEY, J. S. 1877. Catocalae taken at sugar at Center, N.Y. Canad. Entomol., 9:215. 

BARNES, W. & J. MCDUNNOUGH. 1918. Illustrations of the North American Species of 
the Genus Catocala. Mem. Amer. Mus. Natural Hist. new series, vol. 3: part 1, 47 
pp., 22 plates. 

BROWER, A. E. 1974. A list of the Lepidoptera of Maine: part 1, The Macrolepidoptera. 
Life Sci. and Agric. Exp. Sta., Univ. Maine, Orono. Tech. Bull. 66, 136 pp. 

1976. New Catocala of North America (Noctuidae). J. Lepid. Soc., 30:33-37. 

CRUMB, S. E. 1956. The larvae of the Phalaenidae. USDA Tech. Bul. 1135, 356 pp. 

FORBES, W. T. M. 1954. Lepidoptera of New York and Neighboring States: part III, 
Noctuidae. Comell Univ. Agric. Exp. Sta. 433 pp. 

KIMBALL, C. P. 1965. The Lepidoptera of Florida: An Annotated Checklist. Fla. Dept. 
Agric., Div. Plant Industry. 363 pp. 

LINTNER, J. A. 1876. On Catocala pretiosa, n.s. Canad. Entomol., 8:121-122. 

McCaBgE, T. L. & L. M. JOHNSON. Catalogue of the Types in the New York State 
Museum Insect Collection. N.Y.S.M., Albany, N.Y. Bul. 434, 8 pp. 

SARGENT, T. D. 1976. Legion of Night: The Underwing Moths. Univ. Mass. Press, 
Amherst, Mass. xiii + 222 pp., 8 plates. 

SAUNDERS, W. 1876. Notes on Catocalas. Canad. Entomol., 8:72-75. 

SMITH, J. B. 1899. Insects of New Jersey. Supplement to 27th Ann. Rept. State Board 
Agric., Trenton, N.J. 755 pp. 

1910. The Insects of New Jersey. Report of the New Jersey State Museum for 

1909, Trenton, N.J. 888 pp. 


Journal of the Lepidopterists’ Society 
36(1), 1982, 31-41 


AGGREGATION BEHAVIOR IN BALTIMORE CHECKERSPOT 
CATERPILLARS, EUPHYDRYAS PHAETON 
(NYMPHALIDAE) 


NANCY E. STAMP* 
Department of Zoology, University of Maryland, College Park, Maryland 20742 


ABSTRACT. The aggregation behavior of early instars, overwintering fourth in- 
stars, and late instars of Euphydryas phaeton Drury was examined in natural popula- 
tions. Activity outside the webs increased as larvae progressed through the first three 
instars. Mortality at diapause in late summer was 46 to 64% per web. The mean size 
of overwintering larval groups was one-fifth that of mean group size at diapause, as a 
consequence of mortality and group subdivision and separation. Mortality of overwin- 
tering larvae was 18 to 53%. Post-diapause instars were gregarious, until just prior to 
pupation. 


The size of a group of animals can influence the fitness of an indi- 
vidual for a variety of reasons, such as improved thermoregulation, 
feeding facilitation, and effective defensive mechanisms (Allee et al., 
1949; Alexander, 1974; Wilson, 1975; Morse, 1977; Bertram, 1978; 
Stamp, 1980a). In contrast to the larvae of most butterflies, the cat- 
erpillars of the Baltimore checkerspot aggregate in all instars. These 
larvae are conspicuous with their communal webs in mid-summer 
and their coloration and aggregated behavior in the spring. Informa- 
tion on larval aggregation behavior of E. phaeton has been largely 
descriptive. My objectives were to quantify the larval aggregation 
tendencies of early instars, overwintering fourth instars, and late in- 
stars of E. phaeton and to identify factors contributing to aggregation 
behavior. 


METHODS 


I observed E. phaeton from 1977 through 1979 at the Conservation 
and Research Center of the National Zoological Park at Front Royal, 
Warren Co., Virginia. Turtlehead (Chelone glabra L.: Scrophularia- 
ceae), the larval host plant, grew there in dense patches in wet mead- 
ows. This host plant is a clonal, perennial; thus, each plant group 
consisted of numerous stalks. 


Eggs and Early Instars 


Egg clusters were located on turtlehead, and these host plant stalks 
were tagged. To determine the mean number of eggs per cluster, I 
collected and counted the eggs of 32 and 35 new clusters in 1977 and 
1978, respectively. 


* Current address: Department of Zoology, University of Florida, Gainesville, Florida 32611. 


oo JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Twenty-one prediapause larval aggregations were observed at 1000, 
1200, 1400 and 1600 h on three warm, sunny days in July to determine 
the numbers of larvae outside the webs. Webs were collected after 
larval diapause in August 1977, 1978, and 1979; 18, 37, and 10 webs, 


respectively. 


Overwintering Fourth Instars 


Fifteen plots 1 m apart were set up in late October 1978 to deter- 
mine group size of overwintering fourth instars, which leave their 
webs in late fall to overwinter in plant litter on the ground (Bowers, 
1978). To facilitate finding larvae, each plot consisted of a 1 x 1 m 
sheet of clear plastic on the ground. An eighth of a bale of hay was 
used for each plot. The hay was similar in amount and texture to the 
plant litter on the ground at this site. A web tied to a stake was placed 
in the center of each plot. To avoid damaging the webs, the number 
of larvae per web was not determined. However, the mean number 
of larvae per web was probably similar to that of the 1978 set of 
dissected webs (n = 37 webs, x = 110 larvae + 11 S.E.). By mid-No- 
vember, after the caterpillars had moved from the webs into the hay, 
a few spines were marked of each caterpillar within a group, using a 
different color of Testor’s enamel paint for each aggregation on a plot 
to monitor exchange of individuals among the groups over the winter. 
Preliminary tests indicated that the paint (primary and secondary 
colors) did not affect larval behavior. Larvae were checked in mid- 
November and mid-March to determine the number of groups, size 
of groups, and dispersal distance from the webs. 

To determine the number of larvae surviving over the winter, I set 
up a second experiment with 30 boxes containing caterpillars in mid- 
October. Each clear plastic box (13 x 25 x 8 cm) had holes in the 
bottom, covered with window screen for drainage. Tanglefoot was 
sprayed on the upper sides of the box to prevent the caterpillars from 
leaving it. Dried grass in the boxes was similar in texture and depth 
to the plant litter on the ground in the study site. Fifteen boxes each 
contained 100 larvae and no cover, and another 15 boxes each had 
100 larvae and a screen cover. Each box was held in place on the 
ground by four stakes. The surviving larvae were counted in mid- 
March. 


Post-diapause Instars 


To determine the aggregation behavior of late instars, three study 
sites were examined weekly from mid-April through mid-July in 1979. 
The number of larvae within one body length of each other (0.5 to 2 


VOLUME 36, NUMBER 1 33 


cm, depending on the instar) and the number of those greater than 
one body length from each other were recorded. 

In June 1978 and 1979 the patches of turtlehead were mapped in 
two study sites. In both sites 3 x 3 m plots were divided into a 
hundred 30 x 30 cm squares. Plant groups were located within these 
squares. To determine the height of the turtlehead which was avail- 
able in May, plant stalks were chosen by placing a rod through a plant 
group until 30 stalks were partitioned, and these stalks were mea- 
sured. Because there were so few healthy plant groups during this 
period, only three plant groups were sampled. 

To examine aggregation tendencies of late instars further, caterpil- 
lars were collected in mid-April and kept in cages (25 x 25 x 76 cm) 
in the laboratory at 23°C, 70% RH, and 16 h of light, approximating 
conditions in late spring. Ten larvae were placed in each cage with 
pots of snapdragon (Antirrhinum sp.: Scrophulariaceae). Snapdragon 
was chosen because turtlehead was rare in the study sites from April 
through May; snapdragon was easy to grow; E. phaeton larvae feed 
on other Scrophulariaceae in addition to turtlehead (Tietz, 1972); and 
they readily ate snapdragon. The first cage contained one plant about 
25 cm in height, a second cage had two such plants, and a third had 
four plants. Five replicates were run. Observations of the larvae began 
17 h after placing them in the cages and they were monitored at half- 
hour intervals for 3 h. Larvae within a body length (less than 1.5 cm) 
of each other were considered as aggregated. After the experiment, 
all of the larvae pupated; thus, it is unlikely that any behaved differ- 
ently during the experiment due to unapparent parasitism. Further- 
more, in the course of host-parasitoid studies (Stamp, 1980b, 1981b), 
I have not obtained tachinid flies, which are the major parasitoids 
emerging in the pupal stage of E. phaeton. If they occur in this E. 
phaeton population, they are rare. 


RESULTS 
Eggs and Early Instars 


The mean number of eggs per cluster was 273.8 (+23.1 S.D.; 95% 
confidence limits for a mean pooled for years). Of 42 aggregations of 
newly-hatched larvae, 95% moved to the top of the host plant stalk 
before feeding on the turtlehead. 

Larvae fed from late June through early August on leaves enclosed 
within and adjacent to their communal webs. Larval activity outside 
the webs varied through the day (two-way ANOVA after square root 
transformation of count data, P < .001; with interaction between days 
and times of day, P < .05; Fig. 1). In July larval activity outside the 


34 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Third instars 
24 July 


40 


20 
Ke) 
(eb) 
E4 
oD 
ao) . 
D Second instars 
3 17 July 
q 40 
{qe} 
a 
(9e) 
— 20 

First instars 
6 July 
10 


nT a eae | 


1000 1200 1400 1600 . 
Hours 


Fic. 1. Mean number of larvae outside webs (n = 21) with + one standard error. 


webs increased as larvae progressed through the first three instars, 
with significantly fewer first instars outside the webs than during sec- 
ond and third instars (two-way ANOVA, P < .001; Newman-Keuls 
multiple range test (Zar, 1974]; Fig. 1). By the third instar, 86% more 
larvae were on the outside of the webs or feeding on adjacent leaves 
during the day relative to the first instars outside the webs. Larvae 
inside the webs, especially near the center, were quiescent or molt- 
ing. 

The larval aggregations collected after diapause were all in the cen- 
ter of their webs, usually on the upper half of plant stalks. The mean 
number of larvae per web at diapause was 110 to 216, with consid- 
erable variation among years (Fig. 2). Larval mortality due to preda- 
tors within webs was probably small. Few potential predators were 


VOLUME 36, NUMBER 1 35 


300 
18 
is 
® 
= 200 10 
© 
rol 
0) of 
fa} 
6. 
My 100 
1977 1978 1979 
Years 


Fic. 2. Mean number of larvae per web at diapause, with + one standard error. The 
numbers of webs are indicated. 


found in the 65 webs collected in 1977-79: two chrysopid larvae, two 
ant colonies, and 27 spiders. 

Mortality of the egg and larval stages up to diapause was at least 
46%, based on a mean of 274 eggs per cluster and a mean of 148 
larvae (+18 S.E.) per web at diapause for 1977-79. This is a conser- 
vative estimate of mortality at this stage for two reasons. First, some 
webs were composed of larvae from more than one egg cluster on a 
plant stalk. For example, for plant stalks with egg clusters in 1979, 
the mean number of clusters per stalk was 1.5 (or 411 eggs; Stamp, 
1980b). Furthermore, entire egg clusters rarely disappeared (Stamp, 
198la). Based on this mean of multiple egg clusters per stalk and 
mean number of diapausing caterpillars per web, mortality of the egg 
and larval stages up to diapause was 64%. Second, larval aggregations 
seldom split up, even when they defoliated their host plant stalk. The 
caterpillars expanded their web down the plant stalk and onto adja- 
cent leaves, remaining together by using silk trails (e.g., Bush, 1969). 
None of the aggregations observed in this study subdivided. Thus, 
mortality up to diapause was in the range of 46 to 64%. 


36 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


10 


fis] Covered (n=1200) 


Uncovered (n=1100) 


loss 


Percentage of larval 


Total Dead Missing 


Fic. 3. Percentage of larval loss from screen-covered and uncovered boxes over the 
winter. Total numbers of larvae are indicated. The numbers of dead larvae per box 
between covered and uncovered boxes were not significantly different (Mann-Whitney 
U test, P > .20). The numbers of missing larvae between covered and uncovered boxes 
were significantly different (Mann-Whitney U test, P < .002). 


Overwintering Fourth Instars 


The mean size of overwintering larval aggregations was one-fifth 
that of mean group size at diapause, which was a consequence of 
group subdivision and mortality. The sizes of overwintering larval 
groups on plots were similar in November and March (mean of 21.4 
larvae + 24.3 S.D. and 17.9 larvae + 26.9 S.D., respectively; two- 
sample t test, P > .50). Larvae were usually found in dry litter or on 
the plastic sheet 5 to 8 cm below the litter surface. In November most 
groups were tightly aggregated (larvae touching each other), but in 
March most were in loose aggregations (larvae within a body length 
of each other). All of the groups had moved between November and 
March. The larval aggregations in March were farther from their webs 
than in November (11.7 cm + 7.9 S.D. and 20.9 cm + 14.2 S.D., 


VOLUME 36, NUMBER 1 i 37 


n= 2004 530 220 182 166 271176 125 56 41 17 10 
100 


larvae 


50 


Percentage of 
aggregated 


April May June July 


group 


Larvae per 


Fic. 4. Aggregation tendencies among post-diapause larvae. Top—line indicates 
the percentage of aggregated larvae, with total larvae shown by numbers. Factors re- 
lated to aggregation patterns are: A. small patches of turtlehead in mid-April, B. no 
turtlehead above ground in two of three areas, C. no turtlehead in two areas and one- 
third of third area reduced to leafless, 15 cm stalks, D. pupation started and turtlehead 
beginning to reappear, E. turtlehead in all areas and first E. phaeton adults flying, and 
F. pupation ended. Bottom—mean number of larvae per group of those aggregated 
individuals, with + one standard error. 


respectively; normal approximation to Mann-Whitney U test, P < .01). 
Some individuals changed groups over the winter. Over half of the 
larvae originally on the plots in November were missing in March. 
In mid-March there were 1.5 groups per plot (+1.9 S.D.) in contrast 
to 2.3 groups per plot (+1.0 S.D.) in mid-November, but this was not 
a significant difference (Mann-Whitney U test, P > .05). The missing 
groups had either moved off the plots (greater than 45 cm from their 
webs) or were dead. It appears that groups lost some individuals, but 
there is no evidence that entire groups died together. 

Up to 18% of the caterpillars were dead and up to 35% were missing 
in the plastic boxes, for a total larval loss of 53%. The numbers of 
dead larvae per box in the covered and uncovered boxes were similar 
(Fig. 3); however, the numbers of missing larvae per box were greater 
in the uncovered boxes than in the covered boxes. Seven boxes were 
broken by deer and, thus, excluded from the experiment. 


38 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 1. Comparison of defoliated plant stalks and solitary caterpillars. Six com- 
pletely defoliated plant groups were sampled in May, with a total of 397 stalks. The 
226 larvae feeding on these stalks were classified as solitary (greater than a body length 
apart) or aggregated. Statistical analysis showed no correlation between number of 
stalks per plant group and the percentage of solitary larvae per plant group (Spearman 
rank correlation, n = 6, P > .50). Thus, at this time larvae occurring singly did not 
appear to be in response to the limited availability of the larval host plant. 


Plant group 
1 2 3 4 5 6 
Leafless plant stalks ol 33 60 82 91 100 
Percentage of larvae which were solitary 26 26 12 36 AT 0 


Post-diapause Instars 


Some 77% of the late instars were aggregated in mid-April (begin- 
ning of the post-diapause period), but only 20% prior to pupation in 
mid-May (Fig. 4). The mean group size of aggregated larvae was 20.5 
(+52.1 S.D.) in mid-April and 5.0 (+7.1 S.D.) three weeks later. In 
both 1978 and 1979 a few small patches of turtlehead occurred in 
each of three study sites by mid-April. Less than 15% of the caterpil- 
lars were on turtlehead at this time. The patches of turtlehead were 
completely defoliated by late instars in two of three sites by mid-May 
in both years. Caterpillars in the two sites with defoliated turtlehead 
then mainly ate rosaceous shrubs about 10 m from the defoliated host 
plants. In the third area turtlehead was reduced by one third by mid- 
May. Many of the plants had few if any leaves, and the stalks were 
mere stubs, often with one or more caterpillars feeding on them. No 
correlation between the number of stalks per plant group and the 
percentage of single larvae on stalks per plant group was evident at 
this time (Table 1). Thus, the late instars were neither aggregated 
(that is, no negative correlation) nor dispersed (no positive correla- 
tion) in response to the limited amount of turtlehead available to 
them. Shortly after pupation began, the turtlehead began to recover 
and by mid-June the mapped areas of turtlehead very closely resem- 
bled the patches mapped the year before. 

In the laboratory larval aggregation was highest when fewer plants 
were available (two-way ANOVA after square root transformation; for 
the number of plants per cage for all late instars aggregated in cages, 
P < .05; for number of plants per cage for larvae aggregated on plants, 
P < .001). The larvae fed actively, and individuals changed aggrega- 
tions frequently. Half of the larvae were aggregated (mean of 4.7 ag- 
gregated larvae per cage). However, most were aggregated on the 
cages rather than on the plants (mean of only 1.6 aggregated larvae 


VOLUME 36, NUMBER | 39 


on plants per cage). Of those caterpillars aggregated on the cages, 
93% were at the top of the cages. Similarly, in the spring fifth and 
sixth instars were aggregated frequently at heights above the vege- 
tation on dead plant stalks, even when turtlehead was readily avail- 


able. 


DISCUSSION 


The small number of first instars on the outside of webs, relative to 
the numbers of second and third instars, was a consequence of how 
these three instars used the webs. Their abilities to defend them- 
selves differed greatly. First instars were cryptically colored with sin- 
gle hairs extending from tubercles on their bodies. However, second 
and third instars had conspicuous reddish-brown bodies with dense, 
black spines projecting at 45 degree angles from tubercles in rows 
across their bodies. In contrast to first instars, third instars successfully 
knocked parasitoids away from them by head-jerking (Stamp, 1981b). 
Also, second and third instars of E. phaeton were probably toxic to 
some vertebrate predators, because larvae that were reared on turtle- 
head were unpalatable to blue jays (Cyanocitta cristata L.: Bowers, 
1980). Thus, second and third instars of E. phaeton may be better 
protected against parasitoids and predators because of their size, 
spines and toxicity and consequently, less dependent on webs for 
protection against their enemies than first instars (Stamp, 1980b). Sim- 
ilar between-instar differences (in terms of body hair and response by 
predators) also occur in tent caterpillars (Malacosoma americanum 
[Fabricius]; Ayre & Hitchon, 1968). 

The scarcity of first instar E. phaeton on the outside of the webs 
may also be a consequence of adequate food remaining within the 
webs for them. They enclosed the top two furled leaves of the stalk 
with silk and fed within the web. In contrast, second and third instars 
consumed leaves at a faster rate than first instars and often fed from 
the outside of the webs. 

In addition, first instars may be more susceptible to low humidity 
than second and third instars. Morris & Fulton (1970) suggested that 
the webs of fall webworms increased microhabitat humidity, an im- 
portant factor maintaining a high feeding rate and shortening the de- 
velopmental period. Second and third instars may leave the web on 
warm days when they are most active to avoid overheating (Morris 
& Fulton, 1970). 

The movement of groups of fourth instars between November and 
March may be an anti-predatory response, probably directed to in- 
vertebrate predators and insectivorous mammals. Larvae which re- 
main aggregated in the course of such movement may enhance the 


40 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


effect of aposematic coloration and unpalatability on most vertebrate 
predators. Maintaining a clumped distribution also reduces the 
chance of being discovered by predators, and being surrounded by 
others provides individuals with less risk of being attacked (Hamilton, 
1971; Taylor, 1976, 1977). 

Field observations and the aggregation tests in the laboratory sug- 
gest that late instars of E. phaeton were intrinsically gregarious, in 
contrast to a statement that they were largely solitary (Klots, 1951). 
They stayed together in April, sometimes even joining other aggre- 
gations and after defoliation of host plants. Solitary sixth instars col- 
lected during the pupal and adult flight periods were invariably those 
attacked by parasitoids earlier (98% of 95 larvae; Stamp, 1981b). Oc- 
casionally, even these parasitized caterpillars occurred in groups of 
two to four, either on top of leaves or between leaves bound by silk. 
A similar pattern was found for sawfly larvae (Neodiprion swainei 
Midd.), in that after defoliating their host plant, larvae migrated 180 
m or more and continued to aggregate; whereas, diseased larvae were 
solitary (Smirnoff, 1960). The aggregated behavior of late instars of E. 
phaeton may be beneficial by enhancing the defensive mechanisms 
discussed above. 

The decline in aggregation tendencies of late instars was correlated 
with food availability, pattern of larval mortality due to predators and 
parasitoids, and pupation. In areas where the host plants were totally 
defoliated and few other acceptable food plants were available, larvae 
became increasingly dispersed with time. If late instars fed primarily 
on other food plants, it might be advantageous for individuals to be 
solitary if the food they ate rendered them palatable. Bowers (1980) 
demonstrated that E. phaeton larvae were palatable to blue jays when 
as latter instars they fed on English plantain (Plantago lanceolata L.). 
Latter instars were reported to feed on this and a variety of other 
plants (Tietz, 1972). Thus, as a consequence of both the quality and 
quantity of available food, the advantages of aggregating in relation 
to predators may frequently change from one instar to the next. 

These caterpillars were actively gregarious through all instars, this 
behavior contributing to an average group size at each larval stage. It 
is clear that the costs and benefits of aggregation changed as group 
size changed and as these animals progressed through the life cycle, 
a consequence of different factors operating on the larval stages. 


ACKNOWLEDGMENTS 


I am grateful to D. H. Morse for supervising this research and criticizing the manu- 
script. I thank R. F. Denno, J. M. Kemper, D. E. Gill and R. S. Fritz for advice and 
comments on the research and manuscript. J. M. Kemper helped with the field work. 


VOLUME 36, NUMBER 1 4] 


The computer time was supported by the Computer Science Center of the University 
of Maryland. The research was funded by Xerces Society, two Grants-in-Aid from Sigma 
Xi, and the Sigma Xi Chapter of the University of Maryland. I thank the Conservation 
and Research Center of the National Zoological Park at Front Royal, Virginia for use 
of the study area and living accommodations. I am grateful to Grace Russell for typing 
the manuscript. 


LITERATURE CITED 


ALEXANDER, R. D. 1974. The evolution of social behavior. Ann. Rev. Ecol. Syst., 
5:325-383. 

ALLEE, W., O. PARK, A. EMERSON, T. PARK & K. SCHMIDT. 1949. Principles of Animal 
Ecology. Saunders, Philadelphia. 837 pp. 

AyzeE, G. L. & D. E. HircHON. 1968. The predation of tent caterpillars Malacosoma 
americana (Lepidoptera: Lasiocampidae) by ants. Can. Entomol., 100:823-826. 

BERTRAM, B. C. R. 1978. Living in groups: predators and prey. In J. R. Krebs and N. 
B. Davies, eds., Behavioural Ecology, pp. 64-96, Sinauer, Sunderland, Mass. 

BowERs, M. D. 1978. Over-wintering behavior in Euphydryas phaeton (Nymphali- 
dae). J. Lepid. Soc., 32:282-288. 

1980. Unpalatability as a defense strategy of Euphydryas phaeton (Lepidoptera: 
Nymphalidae). Evolution, 34:586-600. 

BusuH, G. L. 1969. Trail laying by larvae of Chlosyne lacinia. Ann. Entomol. Soc. 
Amer., 62:674-675. 

HAMILTON, W. D. 1971. Geometry for the selfish herd. J. Theor. Biol., 31:295-311. 

Kuots, A. 1951. A Field Guide to the Butterflies of North America, East of the Great 
Plains. Houghton Mifflin, Boston. 340 pp. 

Morris, R. F. & W. C. FULTON. 1970. Models for the development and survival of 
Hyphantria cunea in relation to temperature and humidity. Mem. Entomol. Soc. 
Canada No. 70. 

Morse, D. H. 1977. Feeding behavior and predator avoidance in heterospecific 
groups. Bioscience, 27:332-339. 

SMIRNOFF, W. A. 1960. Observations on the migration of larvae of Neodiprion swainei 
Midd. (Hymenoptera: Tenthredinidae). Can. Entomol., 92:957-958. 

STAMP, N. E. 1980a. Egg deposition patterns in butterflies: why do some species 
cluster their eggs rather than deposit them singly? Amer. Natur., 115:367-380. 
1980b. Effect of group size on an egg-clustering butterfly. Ph.D. Dissertation. 

Univ. of Maryland, College Park. 94 pp. 

198la. Parasitism of single and multiple egg clusters of Euphydryas phaeton 

(Nymphalidae). J. N.Y. Entomol. Soc., 89:89-97. 

1981b. Behavioral interactions of parasitoids and Baltimore checkerspot cater- 
pillars (Euphydryas phaeton). Environ. Entomol., 10:in press. 

TAYLOR, R. J. 1976. Value of clumping to prey and the evolutionary response of am- 
bush predators. Amer. Natur., 110:13-29. 

1977. The value of clumping to prey: experiments with a mammalian predator. 
Oecologia, 30:285-294. 

TirETZ, H. M. 1972. An Index to the Described Life Histories, Early Stages and Hosts 
of the Macrolepidoptera of the Continental United States and Canada, Vol. 1. A. 
C. Allyn, Sarasota, Florida. 536 pp. 

WILSON, E. O. 1975. Sociobiology. Belknap, Cambridge, Mass. 697 pp. 

ZAR, J. H. 1974. Biostatistical Analysis. Prentice-Hall, Englewood Cliffs, N. J. 620 pp. 


Journal of the Lepidopterists’ Society 
36(1), 1982, 42-53 


STUDIES ON THE CATOCALA (NOCTUIDAE) OF 
SOUTHERN NEW ENGLAND. VI. THE 
“PAIRING OF C. NEOGAMA AND 
C. RETECTA 


THEODORE D. SARGENT 
Department of Zoology, University of Massachusetts, Amherst, Massachusetts 01003 


ABSTRACT. The closely related, Juglandaceae-feeding Catocala species, C. neo- 
gama and C. retecta, have occurred in nearly equal numbers at the same time of year 
over several years at a single location in southern New England. These two species 
have very similar cryptic forewings, but their hindwings differ markedly. Prior studies 
have suggested that Catocala hindwings function as startle devices in instances of 
avian attack, and that specific differences in hindwing patterns between otherwise 
similar species serve an anti-predator function by interfering with avian habituation to 
startle stimuli. 

Rearing studies with C. neogama and C. retecta indicated that the two species have 
no effects on one another's development rate or survival, though larvae of the two 
species have a somewhat stronger tendency to disperse from one another than do larvae 
of either species alone. Detailed analyses of light-trap data from Washington, Con- 
necticut showed that C. neogama and C. retecta have occurred in equal numbers both 
across and within seasons at that location. However, the variance in the ratios between 
these species over years was greatest both early and late in the season when moth 
abundances were lowest, and this finding is interpreted as evidence for frequency 
dependent selection by birds on this species pair. 


At least 40 species of the noctuid genus Catocala may co-exist at 
a single location in the northeastern United States. A number of these 
species have very similar life histories, including common foodplants 
and nearly identical flight seasons. In some cases, pairs of these 
species have similar bark-like cryptic forewings and resting behaviors 
on tree trunks (Sargent, 1978). 

One of the closest pairings of this sort involves C. neogama (Smith 
& Abbot) and C. retecta Grote (Fig. 1), two closely related Juglan- 
daceae-feeding species (Barnes & McDunnough, 1918; Forbes, 1954). 
These moths were sampled at a light-trap for 12 years by Sidney A. 
Hessel in Washington, Connecticut, and there the two species oc- 
curred in nearly equa! numbers, exhibited parallel fluctuations in an- 
nual abundance, and had essentially identical flight seasons (Fig. 2). 
The two species also have very similar cryptic forewings, though they 
differ markedly with respect to their hindwings—C. neogama having 
orange and black banded hindwings, and C. retecta having entirely 
black hindwings with a prominent white fringe (Fig. 1). 

Catocala hindwings are hidden beneath the cryptic forewings 
when the moths are at rest, and these colorful or boldly patterned 
structures apparently function as startle devices in instances of avian 
attack. The crisp beak-imprints found on the wings of many wild- 


VOLUME 36, NUMBER l 43 


Fic. 1. Catocala neogama (Smith & Abbot) (above) and C. retecta Grote (below). 
Life-size. 


caught specimens provide evidence for this startle function, indicat- 
ing that birds sometimes release these moths when the hindwings are 
exposed (Sargent, 1973). There is also evidence that birds will habit- 
uate, i.e., learn to respond, to particular hindwing patterns (Sargent, 
1973), and I have proposed that the interspecific hindwing diversity 
among the Catocala serves to interfere with this habituation process 
(Sargent, 1969, 1976). This apparent advantage of hindwing diversity 
might then account for the close co-existence of species with very 
different hindwings (Sargent, 1978, 1981). 

Habituation may be defined as the “waning of a response as a result 
of repeated stimulation which is not followed by any kind of rein- 
forcement’ (Thorpe, 1963). This learning process requires a series of 
encounters with the specific stimulus in question, and while some 
generalization to similar stimuli may occur, encounters with suffi- 
ciently different stimuli are known to interfere with, or abolish, the 


44 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


a0 e retecta 404 
° neogama 446 


RaRCENY Ol SRPECiHeES yOUceS 


YEARS 
40 
30 
20 
10 
| 2 3 4 5 6 7 8 


SUCCESSIVE HALF- MONTH PERIODS 


Fic. 2. The annual (above) and seasonal (below) fluctuations in abundance of C. 
neogama and C. retecta at Washington, Connecticut, based on specimens taken in a 
light-trap over 12 years. Abundances are expressed as percentages of the total number 
of individuals taken over the entire 12 seasons for each species (these totals given after 
the species names). The season half-months begin 1-15 July (1) and end 16-31 October 
(8). 


development of an habituated response (Donahoe & Wessells, 1979). 
In the case of predators on C. neogama and C. retecta, it is assumed 
that habituation to the hindwings proceeds so long as only one of the 
two species is being encountered, and that under these circumstances 
the startle response eventually disappears, enabling a predator to cap- 
ture individuals of that species. The number of successive encounters 
required for successful capture of one species, N,, undoubtedly varies 
with many factors, including the species of predator and the time 
elapsing between successive encounters. However, evidence from 


VOLUME 36, NUMBER 1 : AS 


prior studies of avian startle (Blest, 1957; Coppinger, 1969, 1970) sug- 
gests that this value lies somewhere between two and perhaps five or 
six encounters. The presence of a second species with very different 
hindwings serves to reduce the probability of N,. occurring by inter- 
spersing experiences which cause the startle response to reappear. 
Thus, by interfering with the habituation process, both species ben- 
efit by occurring together. 

The extent to which N. would be realized in sampling from a ran- 
domly mixed population of C. neogama and C. retecta would be a 
function of the frequencies of these two species. If N,. was 4, and C. 
neogama and C. retecta occurred in equal numbers (50:50), then the 
probability of a predator encountering four successive individuals of 
one or the other species would be ca. 0.06. The results of similar cal- 
culations, for values of N. between 1 and 5, and for frequencies of 
either species ranging from zero to one, are depicted in Fig. 3. Ex- 
amination of this figure reveals that for values of N, above 3, there is 
a considerable advantage for either species when it is less common 
than the other species. This advantage remains substantial when the 
species are about equally common, but rapidly declines for either 
species as it becomes much more common than the other species. 

These calculations are based on the assumption that predators can- 
not distinguish C. neogama and C. retecta in the resting (cryptic) 
state. If such a distinction were possible, then predators might come 
to associate the two distinguishable cryptic prey with their two very 
different startle patterns and eventually habituate to both. However, 
the forewing similarities of C. neogama and C. retecta may preclude 
the formation of such forewing-hindwing associations and so create 
a very difficult habituation problem. This possibility would provide 
a selective basis for convergence between the two species in cryptic 
characteristics. 

Thus far we have considered only a possible advantage of the co- 
existence of C. neogama and C. retecta. However, these species do 
utilize the same foodplants and one might envision a disadvantage of 
their co-existence with respect to competition for food. I have sug- 
gested previously, however, that Catocala, particularly species like 
these whose larvae feed on the mature green leaves of large decidu- 
ous trees, may not be foodplant limited (Sargent, 1978, 1980). If this 
were true, and if, as previously described, each species benefited with 
respect to predation as the other species increased in abundance (Fig. 
3), then there would be little basis for competitive interference be- 
tween them, and the two species might come to share what is essen- 
tially a single ecological niche. 

The first section of this report is an attempt to shed some light on 


46 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


1.00 
.90 
.80 
5A) 
.60 
.00 
40 
.30 
.20 
10 
.O = 


PROBABILITY OF N, SUCCESSIVE ENCOUNTERS 
JONASNOAIS YSHLO AWOS JO ALINIEVEOud 


0 Ale) ale) Oe tO) LO) 760 10 80 90 1.00 


FREQUENCY 


Fic. 3. The probability of a predator encountering the number of successive indi- 
viduals of one species (e.g., C. neogama) necessary for startle habituation (N,), when 
taking prey from a randomly mixed, two-species population (e.g., C. neogama and C. 
retecta) for various frequencies of the species in question (here C. neogama). Curves 
are plotted for values of N,. from 1 through 5. The probability of an encounter sequence 
other than successive individuals of the species (here C. neogama) is given on the 
right-hand ordinate. 


possible competitive interactions between the larvae of C. neogama 
and C. retecta. These rearing studies were conducted in order to (1) 
corroborate, if possible, the identical phenologies reported from the 
field, (2) compare the survival and development rates of the two 
species when reared in pairs comprised of one or both species, and 
(3) describe any changes in larval behavior associated with the two- 
species as opposed to the one-species rearing condition. 

The second section of this report is devoted to a detailed analysis 
of Hessel’s field data, specifically to determine the extent to which 
C. neogama and C. retecta occurred in equal numbers over the season 
for the 12 years, 1961-1965 and 1967-1971, at Washington, Connect- 


icut. 


A7 


VOLUME 36, NUMBER | 


yt! 


ID 


ray ih Ls 
WWD) 


LLY = 


Deis 


S 


Fic. 4. The rearing situation for C. neogama and C. retecta larvae. Each container 
was supplied with leaves of the foodplant and a twig on which the larva(e) could rest 


(cover of the container not shown). 


Finally, I will discuss the implications of the ecological correspon- 
dence of C. neogama and C. retecta, and speculate as to how their 


numerical equality might be maintained. 
REARING STUDIES 
Methods 


One female each of C. neogama and C. retecta, wild-caught in 
Leverett, Franklin Co., Massachusetts during the summer of 1976, 
were induced to lay eggs in brown paper bags left hanging outdoors. 
The eggs were transferred to small glass jars with perforated lids and 
left in a sheltered, outdoor location to overwinter. Hatching of all the 
eggs occurred on 12 May 1977 for both species. The young larvae 
were immediately provided with leaves of shagbark hickory (Carya 


48 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


—-nmnwfFf OMANDWOO 


retecta x reared with other species 


[_] neogama 


NO. ADULTS ECLOSED 


DAYS 


Fic. 5. The numbers of adult C. neogama and C. retecta eclosing over days (12-20 
July) from larvae reared in single- and mixed-species pairs. 


ovata) for feeding. On 24 May, the larvae were transferred from the 
glass jars and placed in pairs into pint-size plastic containers. Five 
containers with two C. neogama larvae, five containers with two C. 
retecta larvae, and 10 containers with one larva of each species were 
established. Each container was provided with a single hickory twig 
upon which the larvae could rest (Fig. 4). Fresh foodplant was sup- 
plied, and frass removed, on a daily basis. 

The larvae of C. neogama and C. retecta, like those of most Catoc- 
ala, appear very bark-like from the third instar onward and usually 
rest by day on twigs of the hostplant (Sargent, 1976). For 21 days (26 
May-16 June) during this period, the resting positions of the larvae 
in all of the containers having two larvae (i.e., in which neither larvae 
had died or pupated) were noted on one occasion, usually between 
0700 and 0900 h. Data were recorded as to whether both, one, or 
neither larva(e) were on the twig in each container, with the species 
noted in those cases where both species were housed in the same 
container. Records were also kept of larval mortality and the dates of 
pupation and adult eclosion. 


Results 


The phenologies of C. neogama and C. retecta were essentially 
identical under these rearing conditions. The eggs of both species 
hatched on the same date (12 May) and their pupation periods com- 
menced on the same date (9 June). The last pupa of C. neogama was 
formed on 16 June, and the last pupa of C. retecta was formed on 17 
June. The adult eclosion dates overlapped completely, whether the 
moths were reared in single or mixed species pairs (Fig. 5). Thus, the 


VOLUME 36, NUMBER Il 49 


retecta/retecta neogama /neogama retecta /neogama 
122) 
= 70 ] 
Oo 
E 60 4 
& | 
0 1 
@ | 
aro | 
ire 
oO 30 1 7 

— 

5 AO 
lJ 
ro) 
x 10 
uJ 
oa 

2 | ) 2 | fo) 2 | fe) 


NUMBER OF LARVAE ON TWIG 


Fic. 6. The percentage of total daily observations when two, one, or no larvae were 
on the twig in rearing containers housing single-species (retecta/retecta; neogama!/ 
neogama) or mixed-species (retecta/neogama) pairs of larvae. 


species were nearly perfectly synchronized and neither exerted an 
inhibitory effect on the development rate of the other. 

All mortality occurred in the larval stage, but was restricted to C. 
neogama. This mortality was identical, however, in the single species 
and mixed species pairs (50% in both cases). Some aspect of the rear- 
ing conditions was less satisfactory for C. neogama than for C. retecta, 
but this difference was not related to the species with which a larva 
was reared. 

The larvae of the two species exhibited rather similar resting be- 
haviors in the single-species containers (Fig. 6). Both species pre- 
ferred the twig (76.9% of the total observations in C. neogama; 71.7% 
in C. retecta), and the larvae often rested together on the twig (60.0% 
of the observed occasions when at least one larva was on the twig in 
C. neogama; 46.1% in C. retecta). Chi-square contingency tests in- 
dicate that the two species did not differ in their tendencies to utilize 
the twig (x71) = 1.09, P > .30), but that the C. neogama larvae had a 
greater tendency to occur together on the twig (y7,,) = 5.44, P < .02). 

In the mixed-species pairs, both species exhibited a reduced ten- 
dency to utilize the twig (62.0% of the total observations in C. neo- 
gama; 61.4% in C. retecta), and a reduced tendency to occur together 
on the twig (45.5% of the observed occasions when at least one larva 
was on the twig) (Fig. 6). Chi-square contingency tests indicate, how- 
ever, that these changes in behavior from the single species situation 
were only significant for C. neogama (7,1); > 6.6, P < .01). 

The interspecific interactions seemed to involve mild aversion to 
one another's presence, particularly on the part of C. neogama to C. 
retecta. Neither species was clearly dominant on the twig, as the 


50 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


occasions when only C. neogama or only C. retecta was on the twig 
were equally frequent (Fig. 6). There was no evidence of increased 
aggression between species as compared to within species, though 
detailed behavioral observations were not carried out. However, 
whatever the interspecific behavioral interactions in this situation, 
they had no discernible effects on survival or development rates 
(Fig. 5). 


FIELD DATA 
Methods 


The late Sidney A. Hessel of Washington, Connecticut recorded 
the number of individuals of all Catocala species taken in a mercury 
vapor light-trap on virtually every night of every season from 
1961-1965 and 1967-1973. Details of the method employed, and 
many of the data gathered, have been presented elsewhere (Sargent 
& Hessel, 1970; Sargent, 1976, 1977). Here I will focus on details 
relating to the seasonal occurrence of C. neogama and C. retecta at 
this location. 


Results 


As previously reported, C. neogama and C. retecta were taken in 
nearly equal numbers in each of the 12 years of Hessel’s sampling, 
and the two species also showed nearly identical flight seasons (Fig. 
2; Sargent, 1978). Further analysis reveals that the 50:50 proportion 
of C. neogama and C. retecta also obtained across the flight season 
when the data are summed over years (Fig. 7). There was no tendency 
for increasingly close approximation to the 50:50 proportion as the 
flight season progressed. However, the variance in the C. neogama/ 
C. retecta proportions was greatest early and late in the season when 
the numbers of individuals taken were lowest, while this variance 
was least during mid-season when moth abundance was highest. 


DISCUSSION 


The co-existence of C. neogama and C. retecta in nearly equal 
numbers should confer a substantial advantage to both species with 
respect to deterring predators that tend to habituate to specific startle 
patterns (Fig. 3). And we have seen that these species may actually 
occur in nearly equal numbers over many years at a single location 
(Figs. 2 and 7). The question then is, what mechanism might maintain 
this seemingly advantageous numerical relationship of the two 
species? 

The facts that C. neogama and C. retecta are closely related species 
that utilize the same foodplants at the same times suggest that they 


VOLUME 36, NUMBER 1 51 


% ACHROMATIC (refecta) 


| 2 3 4 5 6 7 8 9 10 I 


SUCCESSIVE CALENDAR WEEKS 


Fic. 7. The percentages of C. neogama and C. retecta totals (N) made up by C. 
retecta during successive weeks of the season (30 July—14 October) at Washington, 
Connecticut (solid dots), and the standard deviation for each point (open dots). Data 
are summed over 12 years, 1961-1965 and 1967-1973. 


may be exposed to very similar selection pressures. These same facts 
also suggest that the two species are intense competitors for food. 
However, the results of the rearing studies reported here lend some 
support to prior suggestions that Catocala are not food-plant limited. 
When the two species were reared together, there were some changes 
in larval behavior (Fig. 6), but these seemed to reflect a mild inter- 
specific aversion that resulted in an increased tendency of the larvae 
to disperse. There was no evidence of detrimental aggressive inter- 
actions under these conditions, as neither species exerted an inhibi- 
tory effect on the survival or development rate of the other (Fig. 5). 

Even if one were to grant, however, that food is not limiting for 
these species, it does not follow that they should occur in nearly equal 
numbers over long periods of time. Other factors (e.g., diseases, par- 
asites, climatic variables) would almost certainly favor one or the oth- 
er species at different times. Thus, the extremely close and continu- 
ous 50:50 proportion between C. neogama and C. retecta at 
Washington, Connecticut (Figs. 2 and 7) suggests the operation of a 
rather precise regulatory mechanism. 

A number of mechanisms that are known to promote stable poly- 
morphisms within species (e.g., heterosis and certain types of non- 
random mating) (see Sheppard, 1959) could not operate in this two- 
species situation. However, one extrinsic mechanism, frequency 
dependent selection, could operate here and warrants closer exami- 
nation. 


52 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Frequency dependent selection, or “apostatic’ selection (Clarke, 
1962), occurs when each member of a pair of morphs or species is at 
an advantage with respect to predation when rarer than the other, but 
at a disadvantage when more common than the other (see Cain & 
Sheppard, 1954; Haldane, 1955; Clarke & O'Donald, 1964). Such a 
situation would seem to hold in the present case (see Fig. 3). Thus, 
if C. neogama, for example, comprised 25% of a C. neogamalC. re- 
tecta population at some point in time, then C. retecta should be 
more heavily predated until its numbers were roughly equal to those 
of C. neogama. If C. neogama became more common than C. retecta 
then C. neogama should receive more predation, and so on. In this 
way, predator selection would tend to promote and maintain a 50:50 
proportion of the two species. 

There is some evidence suggesting the operation of frequency de- 
pendent selection in the C. neogama/C. retecta situation at Washing- 
ton, Connecticut. If one assumes that there is some threshold abun- 
dance of these moths below which they do not attract significant 
predation, and above which predation is significant (i.e., that this 
predator-prey relationship fits the so-called Type III function) (Holl- 
ing, 1959, 1965, 1966), then the effects of predation should be maximal 
at mid-season when prey density is highest, and minimal or non-ex- 
istent at the extremes of the season when prey densities are lowest. 
The C. neogamal/C. retecta proportions at Washington, while aver- 
aging 50:50 across the entire season when the data are summed over 
years, do show less variance at mid-season (Fig. 7). This finding may 
be evidence that predators drive the system to equilibrium when the 
moths are abundant, but exert little influence when the moths are 
scarce. And this, in turn, could explain why C. neogama and C. re- 
tecta may not show the 50:50 relationship at locations where they are 
relatively rare (e.g., Leverett, Massachusetts; unpubl. data). 

In summary, there is substantial evidence that C. neogama and C. 
retecta may occur in nearly equal numbers at a given location while 
occupying very similar ecological niches. This situation is seen as a 
consequence of several factors, including (1) a presumed close phy- 
logenetic relationship between the two species, (2) an abundance of 
resources such that food is not limiting, and (3) predator selection that 
(a) promotes convergence in all characteristics associated with cryp- 
sis, (b) simultaneously promotes divergence in startle characteristics, 
and (c) acts in a frequency dependent manner with respect to these 
startle characteristics. 


ACKNOWLEDGMENTS 


The late Sidney A. Hessel generously supplied all of his light-trap data from Wash- 
ington, Connecticut. Early drafts of this paper were constructively criticized by M. 


VOLUME 36, NUMBER 1 53 


Deane Bowers, Alan C. Kamil, and Frederick J. Lindstrom. I extend thanks to M. 
Deane Bowers for assistance in the rearing studies, and to my wife, Katherine, for her 
skillful execution of the figures. 


LITERATURE CITED 


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the genus Catocala. Mem. Amer. Mus. Nat. Hist., 3(1). 47 pp., 22 pl. 

BLEST, A. D. 1957. The function of eye-spot patterns in the Lepidoptera. Behaviour, 
11:209-256. 

CAIN, A. J. & P. M. SHEPPARD. 1954. Natural selection in Cepaea. Genetics, 39:89-116. 

CLARKE, B. 1962. Balanced polymorphism and the diversity of sympatric species, pp. 
47-70. In D. Nichols (ed.), Taxonomy and Geography. Syst. Assoc. Publ. 4. 

& P. O' DONALD. 1964. Frequency-dependent selection. Heredity, 19:201-206. 

COPPINGER, R. P. 1969. The effect of experience and novelty on avian feeding behav- 
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1970. The effect of experience and novelty on avian feeding behavior with 
reference to the evolution of warning coloration in butterflies. II. Reactions of 
naive birds to novel insects. Amer. Nat., 104:323-335. 

DONAHOE, J. W. & M. G. WESSELLS. 1979. Learning, Language and Memory. Harper 
and Row, New York. 513 pp. 

FORBES, W. T. M. 1954. Lepidoptera of New York and Neighboring States. III. Noc- 
tuidae. Cornell Univ. Agric. Exp. Sta. Mem. 329. 433 pp. 

HALDANE, J. B. S. 1955. On the biochemistry of heterosis, and the stabilization of 
polymorphism. Proc. Roy. Soc. London, Ser. B, 144:217—220. 

HOLLING, C. S. 1959. The components of predation as revealed by a study of small- 
mammal predation of the European pine sawfly. Can. Entomol., 91:293-320. 
1965. The functional response of predators to prey density and its role in mim- 

icry and population regulation. Mem. Entomol. Soc. Canada, 45:1-62. 

1966. The functional response of invertebrate predators to prey density. Mem. 
Entomol. Soc. Canada, 48:1-86. 

SARGENT, T. D. 1969. A suggestion regarding hindwing diversity among moths of the 
genus Catocala (Noctuidae). J. Lepid. Soc., 23:261-264. 

1973. Studies on the Catocala (Noctuidae) of southem New England. IV. A 

preliminary analysis of beak-damaged specimens, with discussion of anomaly as 

a potential anti-predator function of hindwing diversity. J. Lepid. Soc., 27:175-192. 

1967. Legion of Night: The Underwing Moths. Univ. Mass. Press, Amherst. 

222 pp: 

1977. Studies on the Catocala (Noctuidae) of southern New England. V. The 

records of Sidney A. Hessel from Washington, Connecticut, 1961-1973. J. Lepid. 

Sacewol- le 15: 

1978. On the maintenance of stability in hindwing diversity among moths of 

the genus Catocala (Lepidoptera: Noctuidae). Evolution, 32:424-434. 

1981. Anti-predator adaptations of underwing moths, pp. 259-284. In A. C. 

Kamil & T. D. Sargent (eds.), Foraging Behavior: Ecological, Ethological, and 

Psychological Approaches. Garland Press, N.Y. 

& S. A. HESSEL. 1970. Studies on the Catocala (Noctuidae) of southern New 
England. I. Abundance and seasonal occurrence of the species, 1961-1969. J. Lep- 
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SHEPPARD, P. M. 1959. Natural Selection and Heredity. Harper & Bro., New York. 
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THORPE, W. H. 1963. Learning and Instinct in Animals, 2nd ed. Harvard Univ. Press, 
Cambridge. 558 pp. 


Journal of the Lepidopterists’ Society 
36(1), 1982, 54-60 


OBSERVATIONS ON THE FLIGHT PERIODICITY OF 
BUTTERFLIES IN WEST MALAYSIA 


A. G. ORR 


School of Australian Environmental Studies, Griffith University, 
Nathan, Queensland 4111 Australia 


ABSTRACT. Data are presented on the flight activity of a large number of butterfly 
species observed over an eight-day period in West Malaysia. There is also discussion 
on variation in the flight periodicity shown by Melanocyma faunula (Westwood) 
(Amathusiidae) across altitude. 


It has often been observed by naturalists who have collected in 
tropical regions that many species of butterflies display a distinctive 
periodicity in their flight activity. However, few empirical data have 
been published to support these observations, and what literature is 
available is sparse and confined to general comments concerning 
broad taxonomic groupings (Corbet & Pendlebury, 1978; Emmel, 
1976) or contained in detailed studies of particular species (for ex- 
ample, Scott, 1974). 


AREA AND METHODS 


During a recent visit to West Malaysia, I collected, over a period 
of eight days, data on the flight activity of a large number of butterfly 
species. The method of data collection was to remain all day at a 
vantage point and note the number of sightings over half-hour inter- 
vals of various species of butterflies. I could record only those species 
with which I was familiar and which were unmistakable in flight; 
hence, I made no records of Lycaenidae or Hesperiidae. 

Data were collected at two sites. The first, Tapah, where the most 
detailed studies were made, was along a steep mountain watercourse 
at an altitude of about 700 m. The vantage point was a rocky outcrop 
beside the stream, and it was from here that I made most of my ob- 
servations. There was a wide break in the forest canopy along the 
stream, so the area received plenty of sunlight from about 0730 h until 
1430 h. The forest was basically of a lowland type, with buttressed 
trees, vines, and large-leafed dipterocarps, although noticeably less 
luxuriant than forest at lower altitudes. : 

The second locality was at Tanah Rata, some 30 km from the first 
site, at an altitude of 1300 m. Here, too, the observation point was 
beside a stream, and the forest was open enough to allow sunlight to 
penetrate for the greater part of the day. The forest was of a stunted 
and open montane type, with smaller leaved trees and no vines. 


VOLUME 36, NUMBER | DO 


In practical terms, an advantage of both sites was that neither af- 
forded too great a field or range of vision, so that I was rarely con- 
fronted with too many butterflies to count or distracted by specimens 
at a distance too great for them to be identified quickly and accurately. 
In addition, overall population numbers at both sites were quite low, 
which permitted an accurate appreciation of butterfly activity at all 
times during the period of observation. 


RESULTS 


The observations are summarized in Tables 1, 2 and 3. For each 
species, numbers of sightings for each hourly interval from 0700 to 
1800 h are expressed as a percentage of total number sighted. At 
Tapah data were collected on five days and are summarized in Table 
1. Data were recorded for three days at Tanah Rata (1300 m). Table 
2, summarizes these data for the first two days, which were both cloudy 
after 1100 h with only very occasional sunshine after 1200 h. Table 
3 contains the data for the third day, which was sunny throughout. 
This division was made because it was noted that M. faunula were 
more active on days when the sky was overcast. The dates, and details 
of collecting times, are listed below each table. 

Six species have been selected to illustrate various patterns of flight 
activity at Tapah, and histograms of percentage of sightings against 
time of sighting are given in Fig. 1. Fig. 2 illustrates the changing 
pattern of M. faunula activity with altitude and weather conditions. 


DISCUSSION 


Although the majority of butterflies were observed to fly between 
0900 h and 1500 h, there are many which do not conform to this 
pattern. 

Most Papilionidae, Pieridae, Danaidae and Nymphalidae are strict- 
ly diurnal, but there is a wide range of variation in duration of flight 
period and time of maximal activity. 

Certain species remain active throughout the day but reserve par- 
ticular periods for the different life functions. The Neotropical ith- 
omiid, Mechanitis isthmia, visits flowers in the early morning and 
late afternoon, and devotes the warmer part of the day to courtship, 
mating, and oviposition, activities which take place in the shade (Em- 
mel, 1976). My own observations in Malaysia indicate that many 
species of Euthalia (Nymphalidae) will fly around the sunlit forest 
margins in the early morning but move into the deep jungle later in 
the day. 

The histograms for Trogonoptera and Graphium (Figs. la and 1b) 
show the type of pattern typical for most Papilionidae, Pieridae, Da- 


56 


TABLE 1. Observed flight activity of western Malaysian butterflies. 


PAPILIONIDAE 


Trogonoptera brookiana 3 


T. brookiana 2 

T. amphrysus 
Atrophenura varuna 
A. sycorax 

Papilio helenus 

P. iswara 

P. nephelus 
Graphium evemon 
G. antiphates 

G. macareus 


PIERIDAE 


Leptosa nina 
Cepora nadina 
Appias lyncida 
A. indra 

A. nero 
Pareronia valeria 
*Eurema spp. 
Gandaca harina 


DANAIDAE 


Danaus aspasia 
Ideopsis gaura 
Euploea mulciber 
E. camaralzeman 
E. diocletianus 


SATYRIDAE 


*Ypthima fasciata 
*Y. pandocus 
Ragadia crisilda 
Melanitis leda 


AMATHUSIIDAE 


Amathusia spp. 
Faunis gracilis 
Melanocyma faunula 
Xanthotaenia busiris 
Thauria aliris 


NYMPHALIDAE 


Cupha erymanthis 
Terinos terpander 
Cyrestis nivea 
Parthenos sylvia 
Stibochiona nicea 
Polyura spp. 
Charaxes bernardus 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


700 800 


3.9 
6.7 


3.6 
Wy 


1.3 


0.9 


900 


14.8 


143 


20.7 


1000 


Syl 


14.7 
20.3 
5.6 


1100 


3.1 


10.5 


* Figures indicate captures rather than sightings. Dates and times of observations: 5-I-79 700-1800 h, 7-I-79 
600-1900 h, 9-I-79 700-1900 h, 10-I-79 600-1800 h, 13-I-79 700-1800 h. 


VOLUME 36, NUMBER l1 


1200 


29.4 
21.1 
27.8 
30.0 


47.4 
46.7 


1300 


6.3 


IS 
U7. 
22.2 
30.0 


26.3 
26.7 


1400 


TABLE 1. Continued. 


6.3 


1500 


2.0 


5.3 


2.3 


9.8 


1600 


3.6 


3.8 


Dail 
9.8 


33.3 


14.3 
26.7 
28.1 
33.3 


3.5 


1700 


0.7 


Well 


1.3 


1800 


Total 


57 


58 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 2. Observed flight activity of select species. 


800 900 1000 1100 1200 1300 1400 1500 1600 1700 1800 Total 


Papilio helenus 12.5 50.0 25.0 125) 8 


Delias descombesi 25.0 45.0 20.0 10.0 20 
Melanocyma 
faunula 98 75 12:3-4179 17.0 TS 13:2 855 ese cela 


Dates and time of observations: 8-I-79 800-1800 h, 11-I-79 800-1800 h. 


naidae, and many Nymphalidae and Satyridae. The flight period may 
be more concentrated as in Graphium, or spaced out as in Trogon- 
optera, but in both cases more activity occurs between 1000 h and 
1400 h. Unlike the other papilionids, Atropheneura display a less 
regular flight pattern (possibly a function of the small sample size), 
but distinctly avoid the hotter parts of the day (Fig. lc). 

The Charaxinae (Fig. 1d) have a brief period of activity in the early 
afternoon, with over 90% of observations being made between 1200 
and 1500 h. By contrast, Gandaca harina (Horsefield) (Fig. le) prefers 
the early part of the day but is on the wing for a longer period. Perhaps 
this can be explained by the very powerful flight of the Charaxinae, 
which when active, pause only briefly to refresh themselves, usually 
at dung or carrion. Gandaca harina flutters weakly, always around 
the same tree, and settles frequently. 

Faunis gracilis (Butler) (Fig. 1f) exhibits a crepuscular flight pattern 
typical of the Amathusiidae. Records were not kept after 1800 h, ow- 
ing to poor light, but a number were seen at this time. The related 
Discophora timora (Westwood) has been recorded at fruit bait as late 
as 2130 h (Corbet & Pendlebury, 1978). In M. faunula, the crepus- 
cular habit is less pronounced, but the bulk of flight activity at 700 m 
is concentrated towards evening (Fig. 2a). At 1300 m it flies through- 
out the day, though it is more active in the afternoon on sunny days 
(Fig. 2c). On cloudy days at 1300 m, there was some reduction in 
activity (Fig. 2b) but not to the same extent as observed in Delias 


TABLE 3. Observed flight activity of select species. 


800 900 1000 1100 1200 1300 1400 1500 1600 1700 1800 Total 


Papilio helenus 12:5 76:35 Aes 1S:8es12%) WSS eles 16 
Delias descombesi S44 S4y 103 13:8) 20 24a 17-9 29 
Melanocyma 

faunula 29° 2:9 8.7 13.0 11.6- 15:9 13:0" 174° One 


Date and time of observation: 15-I-79 800-1800 h. 


VOLUME 36, NUMBER 1 


= nm oO 
fo) ro) ro) 


Oo 


total observations 
s.hUS8lhCUDS 
(er) 


co) 


@ 


Percentage of 
S 


(o>) 


aa 


oe 10° 12 4 i 18 
T Ime («klOOnr) 
Fic. 1. 


59 


b 


30 

20 

10 

0 le 
18 
18 


0 10:4 12> 14- le 
50 
40 
30 d 
20 
10 
0 

0 lo) 2 4 16 
30 

ca Ty 

0 One 14 16 


2 


10 


8 
8 
0 
8 


| 
TiIrmetx!OOnr) 


18 


Histograms illustrating flight activity of various butterfly species. (a) Tro- 


gonoptera brookiana; (b) Graphium spp.; (e) Atrophencura spp.; (d) Polyura and 
Charaxes spp.; (e) Gandaca harina; (f) Faunis gracilis. (Histograms represent pooled 
results for both sexes.) 


descombesi (Boisduval), a montane species not found at Tapah, which 
only took to wing during short periods of sunshine. 

Papilio helenus (Linnaeus) displays basically similar flight patterns 
at 700 and 1300 m. The different flight patterns of Melanocyma at 
different altitudes suggest that temperature may be an important fac- 
tor in determining activity. 


b c 


Percent total obsns. 
S26 “SSeS 
a 
= ie) 
(o>) oO 
Soon ee 


0 
Ogre). 12. lay. 16. 16 OS win lee 14716: 118 08 10" (24 | IGr te 
Time loOnr) Time («*lOOnr) Time (xlOOnr) 


Fic. 2. Histograms illustrating flight activity of Melanocyma faunula under differ- 
ing conditions. (a) 700 m, sunny day; (b) 1300 m, cloudy day; (ce) 1300 m, sunny day. 
(Histograms represent pooled results for both sexes.) 


60 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


ACKNOWLEDGEMENTS 


I would like to thank Dr. R. L. Kitching for reviewing the manuscript. 


LITERATURE CITED 


CoRBET, A. & H. M. PENDLEBURY. 1978. The Butterflies of the Malay Peninsula. 
Third Edition revised by J. N. Eliot. Malayan Nature Society, Kuala Lumpur, 
Malaysia. 

EMMEL, T. C. 1976. Butterflies. Thames and Hudson, London. 

FLEMING, W. A. 1975. Butterflies of West Malaysia and Singapore, Vol. 1. E. W. Clas- 
sey. Faringdon, Berkshire, England. 

Scott, A. 1974. The interaction of behavior, population biology and environment in 
Hypaurotis crysalus (Lepidoptera). Amer. Midl. Nat., 91: 383-394. 


GENERAL NOTE 


Journal of the Lepidopterists’ Society 
36(1), 1982, 60-61 


DOWNY WOODPECKERS AS PREDATORS OF 
HYALOPHORA CECROPIA PUPAE 


The leaf-drop of a large silver maple (Acer saccharinum L.) in my yard during the 
fall of 1979 revealed 12 cocoons of Hyalophora cecropia (Hubner) scattered among its 
branches, many twenty to thirty feet above the ground. These pupae developed from 
eggs laid by the captivity-mated females I released the preceding spring. I felt content 
in thinking there would be a good source of wild males should I need them for back- 
crosses in my hybridization studies in this moth genus. Consequently, I decided not 
to collect them for storage as I usually do. 

On 20 December 1979, as I returned from a late afternoon walk on an unusually 
clear cold day, I heard a pecking noise in the tree as I passed beneath it. To my chagrin, 
ona high branch a female downy woodpecker (Dendrocopos pubescens L.) was pecking 
one of the cocoons. It was perched on the cocoon, pecking with some difficulty as the 
flexible branch moved with the force of its bill. I repeatedly tried to drive it away but 
it always quickly returned and continued to peck. This experience recalled my dis- 
cussion with Drs. Dale F. Schweitzer and Charles L. Remington at Yale University 
earlier in the season of jays feeding on the pupa in cocoons and the studies of Wald- 
bauer, Sternburg et al. (1967, Ann. Entomol. Soc. Amer., 60: 97-101; 1967, Ecology, 
48: 312-315; 1970, Ann. Entomol. Soc. Amer., 63: 1366-1369) in Illinois of woodpecker 
predation on Hyalophora cecropia cocoons. 

I then gathered all the cocoons I could reach with a twelve-foot extension pruner. 
Most had holes in them, as shown in Fig. 1. I opened several to find in most a shriveled 
pupal shell, its contents cleverly removed, likely by the tongue of the woodpecker. Of 
the seven cocoons I was able to reach, only one appeared to contain a living pupa. 

I was further surprised to observe that the woodpecker seems to know which cocoons 
contained viable pupae, since those containing parasitized pupae appeared to be pur- 
posely avoided. Two cocoons contained dead larvae with tell-tale shriveled egg shells 
of tachinid parasites on their surfaces. 


VOLUME 36, NUMBER 1 | 61 


eee ee a ee ura MUU LULA LU Un EARLS EMU Lu RED 
0 10 20 30 40 50 60 70 80 90 


Fic. 1. Cocoon of Hyalophora cecropia with evidence of predation. 


Perhaps, the woodpecker is able to hear or feel the rattle of a dry, shriveled pupae 
in the cocoon as it pecks, and hence avoids the wasted energy of drilling a hole through 
a tough pupal case only to find no food within. 

It was extremely interesting to observe the behavior of this woodpecker as it me- 
thodically destroyed the remaining viable cocoons beyond my reach in the tree. There 
is no doubt woodpeckers find pupal Saturniidae a highly attractive food source during 
the winter. 


THOMAS R. MANLEY, Department of Biology, Bloomsburg State College, Blooms- 
burg, Pennsylvania 17815. 


Journal of the Lepidopterists’ Society 
36(1), 1982, 62-64 


OBITUARY 


WILLIAM DONALD PATTERSON, JR., 1905-1980 


William Donald Patterson, Jr., a sustaining member of the Lepidopterists’ Society 
since 1953, died suddenly at his home in Atherton, California on 3 March 1980. Don’s 
interest in Lepidoptera was avocational. He was interested in field work and in the 
support of studies of the butterfly and skipper fauna of Baja California, Mexico, and he 
further supported field work of students with other entomological projects in this area. 

Don was born on 1 July 1905, in San Francisco, California. The first of three sons of 
William Donald Patterson and May Bird Patterson, he was raised in Centerville (now 
part of Fremont) and attended Piedmont High School. He graduated from Harvard 
University with an A.B. degree in Mining Engineering in 1927 and from Stanford 
University with an M.A. degree from the Graduate School of Business in 1930. In 1938 
he was married to Dorothy Eden Wilcox. 

Don, a third-generation Califomian, a rancher and investor, operated the 3000 acre 
Patterson Ranch near Newark, Alameda Co., as well as a 10,000 acre cattle ranch near 
Livermore. His grandfather, George W. Patterson, came to California from Indiana in 
1849 to join in the search for gold, and shortly thereafter bought a portion of the Spanish 
land grant—Rancho de los Potreros de los Cerritos, which is now a part of the city of 
Fremont. The proximity of the ranch to San Francisco Bay permitted, in earlier days, 
the shipment of livestock and produce (especially wheat) by shallow draft schooners 
from a nearby slough to the port of San Francisco. The Livermore Ranch was acquired 
at a later time and has now become divided by the Lake Del Valle State Recreational 
Area, the establishment of which Don supported. Presently, this is a very popular 
aquatic recreational area; even though, as his son Bill recalls, this once was a beautiful 
valley with a sycamore lined creek. Thomas W. Davies remembers this area as the 
habitat of a large population of Speyeria callippe near comstocki (Gunder), which still 
occurs in smaller numbers on the slopes of the surrounding hills. The ranch is the type 
locality for the glyphiperygid, Choreutis apocynoglossa Heppner (1976, Pan-Pacific 
Entomol., 52(3):256—-262). In the early 1970’s Don was interested in donating a portion 
of the ranch’s oak woodland as a preserve, but unfortunately, there was no institutional 
support available for its establishment. 

Don was an outdoors person, who began to collect butterflies and moths in his teens. 
He acquired some of the Lepidoptera books of his time—by Wright, Holland, Mc- 
Glashan, and others, and he purchased specimens as well from a dealer in Hope, 
Arkansas. These collections became inactive and were stored in an attic. In the early 
1950’s his interest was renewed when his son Bill began collecting butterflies (the boy 
also discovered and salvaged specimens from the stored collections). In 1953 Don 
joined the Lepidopterists’ Society as a sustaining member. 

Don and Bill attended the 1954 Pacific Slope meetings of the society, in San Fran- 
cisco, at the California Academy of Sciences. In 1955, Don was Secretary Pro. Tem. to 
the Second Pacific Slope meetings held at San Diego. At the Santa Barbara meetings 
in 1958 he presented an invited paper on the butterflies of the Sierra San Pedro Martir. 
It was during this period through contact with Dr. Jerry A. Powell, Charles F. Harbin- 
son, and others, that his interest in Baja California butterflies was kindled (strengthened 
by his basic love of the peninsula). In late May and early June 1958, Dr. Powell, Bill 
Patterson, and Don made a 90 mile journey by foot, with mules carrying their gear, 
into the Sierra San Pedro Martir, Baja California Norte. Theirs was the first group of 
lepidopterists to explore the high tablelands with its coniferous forests. An interesting 
account of this exploration and record of species collected was published by Don & 
Powell (1960, J. Lepid. Soc., 13(4):229-235). 

Don participated in over 20 trips into both the northern and southern states of Baja 
Califomia, many of these for the purpose of collecting Lepidoptera. Most of the field 
work was undertaken before the present paved road was built, so that the country was 
rugged to explore. The trips were made by jeep, horseback, or by hiking and using 


VOLUME 36, NUMBER 1 63 


WILLIAM DONALD PATTERSON, JR., 1905-1980. 
(Photography by F. Ramsdell Cummings.) 


pack animals. Trip participants included his wife Dorothy, his son Bill, Dr. Ira L. 
Wiggins (on three trips), Joe Donohoe (on three trips), Dr. Powell, Dr. John T. Doyen, 
Sigurd L. Szerlip, Herman G. Real, and Colonel Forde. Joe Donohoe recalls Don’s 
speciality of making soda biscuits in a reflector oven. Several thousand butterflies and 
skippers were collected as a result of the field work. Don was particularly pleased with 
the discovery and the patronym of Erynnis tristis pattersoni Burns (1964, Univ. Calif. 
Publ. Entomol., 37:143), and the co-discovery with Powell of Apodemia mormo di- 
aleuca Opler & Powell (1962, J. Lepid. Soc., 15(3): 167-168). The Erynnis was collected 
in May 1959, in the Sierra de la Victoria, Baja California del Sur, and the Apodemia in 
May 1958, in the Sierra San Pedro Martir. More recently the scorpion, Vaejovis pat- 
tersoni Williams & Haradon, was described (1980, Occ. Pap. Calif. Acad. Sci., 
135:65-66). 

Don was interested in sponsoring research on the skippers and butterflies of Baja 
California, even though he did not anticipate publishing in this field himself. He hoped 
that such studies would contribute to a better understanding of the origin and rela- 
tionships of the fauna of this peninsula. Arrangements were made in May 1969, to 
finance a post-doctoral position at the California Academy of Sciences with his com- 
mitment to provide a stipend for the first year of this study. A candidate for the position 
was selected, but at the last moment, that person decided otherwise. Starting in 1970, 


64 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Herman G. Real, then a student at San Jose State University, received partial support 
for several years, during which time he completed both his A.B. and M.A. degrees. His 
research focused on this region, and a large illustrated manuscript entitled “The Dis- 
tribution of Skippers and Butterflies in Baja California, Mexico,’ was completed. This 
manuscript will be edited for publication in the near future. 

Don also provided “seed money’ to aid students in field work in Baja California. 
Small grants, usually of about $500 each, were awarded through the California Academy 
of Sciences or the Explorers’ Club. Between 1974 and 1979 the following persons 
received such awards (including the year and topic of research): Richard M. Haradon 
(1974, Scorpions); Warren E. Savary (1975, Solpugida); Dr. David B. Weissman (1977 
and 1978, Orthoptera); Eric M. Fisher (1977, Diptera: Asilidae; in 1978, for field work 
at Barro Colorado Island, Panama); and Gay C. Hunter (1979, Diptera: Bombyliidae). 
In 1976 he provided the funds needed by the Academy to purchase the Robert 
G. Wind collection of North American butterflies (made available through the interest 
of Mrs. Clo Carroll). 

Don was a very thoughtful man and an active person with many interests. He served 
as 80th president of the Society of California Pioneers in 1966-67. He served on the 
Board of Directors of the San Francisco Zoological Society and participated in a 1970 
expedition to Nepal to collect an Indian Rhinoceros. This expedition utilized elephant 
transportation, and collections of butterflies were also made at this time. He also served 
on the Boards of Directors of the English Speaking Union of San Francisco and of the 
Leslie Salt Company. Don also held memberships in the Pacific Union Club of San 
Francisco and the Explorers’ Club of New York and London. He was a West Coast 
Director of the San Francisco chapter of the Explorers’ Club. In addition, Don was a 
Field Associate and Fellow (elected in 1971) of the California Academy of Sciences, 
a Vesterman of the Holy Trinity Episcopal Church in Menlo Park, and a member of 
the Atherton Town Planning Commission for 15 years. During the last 10 years, Don 
and his wife Dorothy spent each May in London, where they resided in the Sloane 
Square area. Don was a Fellow of the Royal Geographical Society and a member 
of both the English Speaking Union of London and the Zoological Society of London. 
During the Second World War, he oversaw work at the Joshua Hendy Iron Works 
of Sunnyvale, California. . 

Don is survived by his wife, Dorothy, Atherton, California, and 5.children: William 
D. Patterson, III, Sacramento; Wilcox Patterson, Woodside; George N. Patterson, Palo 
Alto; Mrs. Gerald Green, Ashland, Oregon; and Mrs. Eden Patterson, Jamestown, 
Tennessee; four grandchildren; and a brother, David G. Patterson, Alamo, California. 

I am indebted to Mrs. Dorothy W. Patterson, Mr. William D. Patterson, III, Mr. 
Thomas W. Davies, Mr. Joseph A. Donohoe, IV, Mr. J. Roger Jobson, Dr. Jerry A. 
Powell, and Dr. Ira L. Wiggins for information contained in this article. 


PAUL H. ARNAUD, JR., California Academy of Sciences, Golden Gate Park, San 
Francisco, California 94118. 


Date of Issue (Vol. 36, No. 1): 16 June 1982 


EDITORIAL STAFF OF THE JOURNAL 
THOMAS D. EICHLIN, Editor 


% Insect Taxonomy Laboratory 
1220 N Street 
Sacramento, California 95814 U.S.A. 


J MAGDA R. PAPP, Editorial Assistant 
DOUGLAS C. FERGUSON, Associate Editor THEODORE D. SARGENT, Associate Editor 


NOTICE TO CONTRIBUTORS 


Contributions to the Journal may deal with any aspect of the collection and study of 
Lepidoptera. Contributors should prepare manuscripts according to the following in- 
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Literature Cited: References in the text of articles should be given as, Sheppard 
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the heading LITERATURE CITED, in the following format: 


SHEPPARD, P. M. 1959. Natural selection and heredity. 2nd. ed. Hutchinson, London. 
209 pp. 

196la. Some contributions to population genetics resulting from the study of 
the Lepidoptera. Adv. Genet. 10: 165-216. 


In the case of general notes, references should be given in the text as, Sheppard (1961, 
Adv. Genet. 10: 165-216) or (Sheppard 1961, Sym. R. Entomol. Soc. London 1: 23-30). 

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CONTENTS 


THE BUTTERFLY FAUNA OF BARTON CREEK CANYON ON THE 
BALCONES FAULT.ZONE, AUSTIN, TEXAS, AND A REGIONAL 
List.. Christopher J. Durden =... 


THE LARVA AND STATUS OF CATOCALA PRETIOSA (NOCTUIDAE), 
WITH DESIGNATION OF A LECTOTYPE. Dale F. Schweitzer 


AGGREGATION BEHAVIOR IN BALTIMORE CHECKERSPOT CATER- 
PILLARS, EUPHYDRYAS PHAETON (NYMPHALIDAE). Nancy E. 
Sarpy Ss 


STUDIES ON THE CATOCALA (NOCTUIDAE) OF SOUTHERN NEW 
ENGLAND. VI. THE “PAIRING” OF C. EOGAMA AND C. RETECTA. 
Theodore D. Sargent, 2... 


OBSERVATIONS ON THE FLIGHT PERIODICITY OF BUTTERFLIES 
IN WEST MALAYSIA. A.G. Orr (0000) Ue 


GENERAL NOTE 


Downy woodpeckers as predators of Hyalophora cecropia pupae. Thomas 
R. Manley 


OBITUARY 
William Donald Patterson, Jr., 1905-1980. Paul H. Arnaud, Jr. ---------- 


18 


31 


42 


62 


Volume 36 1982 Number 2 


ISSN 0024-0966 


JOURNAL 


of the 


_ LEPIDOPTERISTS’ SOCIETY 


- 
re 
a 

ve 


x 


Published quarterly by THE LEPIDOPTERISTS’ SOCIETY 


Publié par LA SOCIETE DES LEPIDOPTERISTES 
Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN 
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Pith GOOG 


24 September 1982 — 


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Cover illustration: Mature larva of Eumorpha fasciata Sulzer (Sphingidae) feeding 
on Ludwigia sp. (Jussiaea) in southern Florida, where this hawk moth is generally 
found throughout the year. Original drawing by Mr. John V. Calhoun, 382 Tradewind 
Ct., Westerville, Ohio 43081, USA. 


JOURNAL OF 


Tue LeEPIDOPTERISTS’ SOCIETY 


Volume 36 1982 Number 2 


Journal of the Lepidopterists’ Society 
36(2), 1982, 65-75 


FOODPLANT AND OVIPOSITION RECORDS FOR 
PANAMANIAN LYCAENIDAE AND RIODINIDAE 


ROBERT K. ROBBINS! AND ANNETTE AIELLO 


Smithsonian Tropical Research Institute, Box 2072, 
Balboa, Republic of Panama 


ABSTRACT. We present larval foodplant and female oviposition records for 15 
Panamanian butterfly species in the Lycaenidae and Riodinidae. Many of these species 
feed on reproductive parts of plants, e.g. flowers, rather than foliage. Some species are 
facultatively myrmecophilous, and one species may have an obligate relationship with 
ants. We discuss possible biological consequences of flower-feeding for lycaenid but- 
terflies. 


Larval foodplant records of Lycaenidae and Riodinidae are of par- 
ticular interest for several reasons. First, many of these species feed 
as larvae on the flower-buds, flowers, and fruits of plants (Downey, 
1962), and thus may exert stronger selective forces on their foodplants 
than foliage feeders (e.g. Breedlove & Ehrlich, 1968). Plant responses 
may include changes in flowering phenology (Breedlove & Ehrlich, 
1968; Ehrlich et al., 1972) and synthesis of a variety of secondary 
compounds in flowers (Dolinger et al., 1973). The selective forces 
acting in turn on the butterflies, however, are unclear at present. Sec- 
ond, larvae of these related families utilize as a group a particularly 
broad spectrum of foods, including insect prey (Ehrlich & Raven, 
1964). In addition, some species are unusually polyphagous (Downey, 
1962). And third, the larvae of many lycaenid and riodinid species are 
tended by ants, a putatively mutualistic interaction (e.g. Hinton, 1951; 
Callaghan, 1977). 

Larval foodplant records for Neotropical lycaenids and riodinids are 
more poorly known than those for other biogeographical regions 
(Downey, 1962). Many more records for this species-rich fauna need 
to be accumulated before general patterns of biological interest can 
be deduced or tested. We report here foodplant records that we made 


‘Current address: Department of Entomology, NHB 127, Smithsonian Institution, Washington, D.C. 20560. 


66 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


on Barro Colorado Island (BCI), Panama, a research station adminis- 
tered by the Smithsonian Tropical Research Institute, and in sur- 
rounding areas of Panama Province, Republic of Panama, during the 
past few years. We then discuss some possible consequences of flow- 
er-feeding by larvae and comment on the extent of myrmecophily in 
Panamanian riodinids. 

Identification and deposition of insects and plants are as follows: 
Plants were identified by the junior author, by comparison with spec- 
imens in the Missouri Botanical Garden’s herbarium in Ancon, Pan- 
ama, and by various visiting botanists. Gordon B. Small of Balboa, 
Panama, and the senior author identified the butterflies by compari- 
son with type specimens in the U.S. National Museum, with photo- 
graphs of type specimens in the British Museum (Natural History), 
and with published plates and descriptions. Taxonomic notes are added 
where necessary to clarify identifications. Many of the species do not 
have valid generic names, and are placed for convenience in Thecla 
F., which correctly applies to species in a different tribe (Eliot, 1973). 
Larval head capsules, pupal skins, and adult specimens for species 
denoted by “Aiello lot #” are deposited in the collection of the junior 
author. Adult specimens and some preserved immature stages of other 
species are retained in the collection of the senior author. 

We have had difficulty rearing many of these species from the egg 
with the notable exception of Arawacus aetolus Sulzer. As a result, 
some of our “foodplant’” records are oviposition sightings. Other 
workers may be able to confirm whether or not these plants are suit- 
able larval foodplants. 


Lycaenidae 


Arawacus aetolus lincoides (Draudt): We saw females ovipositing 
on the leaves and twigs of Solanum lancaeifolium in Gamboa on 10 
September 1979, and 6 & 10 December 1979. Unlike many other 
lycaenids, females do not lay eggs on flowers or maturing fruits. Fe- 
males oviposit readily on this plant in the lab, and we have reared 
more than a hundred individuals from eggs on the leaves of S. lan- 
caeifolium. The ithomiine butterflies, Mechanitis lysimia macrinus 
and M. polymnia isthmia also oviposit on this plant species. 

Although third and fourth instar larvae are often tended by the ants 
Ectatomma tuberculatum, E. ruidum, and Pheidole sp., they are reared 
easily in the lab without ants. A final (fourth) instar larva that we found 
in Gamboa on 2 January 1980 was tended by Pheidole and had a 
chalcidid wasp flying around it. This larva pupated between 4 & 6 
January, and a chalcidid emerged on 25 January. 

On 18 January 1980 we watched a female of A. aetolus oviposit on 


VOLUME 36, NUMBER 2 67 


Solanum ochraceo-ferrugineum along the road to Cerro Campana at 
500 m. We found two other eggs and seven larvae on this plant. Two 
males and a female reared on leaves eclosed on 10 & 12 February. 
We switched another larva to S. lancaeifolium after the third molt. It 
pupated on 4 February, and eclosed as a female on 14 February. In 
addition, the female that had oviposited on S. ochraceo-ferrugineum 
laid another 49 eggs on S. lancaeifolium in the lab. Thirty-five of these 
eggs were reared to adult. 

Boyce A. Drummond III reared three specimens of A. aetolus sepa- 
rata Lathy in Limoncocha (Rio Napo), Ecuador, on Solanum cocon- 
illa. A larva that he found on 4 June 1974 pupated on 8 June, and 
eclosed as a male on 19 June. Two larvae that he found on 19 June 
1974 pupated on 26 June, and eclosed on 8 July (female) and 9 July 
(male). 

Guppy (1904) reported the larval foodplant of the nominate sub- 
species from Trinidad as cocoa, but Kaye (1921) corrected the record 
to Solanum sp. It is likely that larvae of A. aetolus feed on a number 
of species of Solanum throughout its range. 


Taxonomic note: Our identification of this species (and its subspecies) is based on 
an unfinished ms. of H. K. Clench. The senior author is completing this paper. 


Tmolus echion (L.): We found a larva on leaves of Stigmaphyllon 
lindenianum (Malpighiaceae) on BCI on 31 May 1978. It pupated on 
16 June, and eclosed on 29 June (Aiello lot 78-67). 

A female of T. echion oviposited on a flower of Aphelandra dep- 
peana (Acanthaceae) among numerous ants (Ectatomma sp.) on Ta- 
boga Island 12 miles off the southern coast of Panama on 24 Decem- 
ber 1978. The ants were feeding on secretions from extra-floral 
nectaries on the flower bracts. We also found two egg shells on this 
flower, plus damage similar to that produced by lycanid larvae. No 
larvae were reared. 

T. echion is of economic importance because it feeds on flowers of 
the weed Lantana camara (Verbenaceae). There are specimens reared 
from L. camara in the United States National Museum from Brasil 
(Sao Paulo, Rio de Janeiro, and Minas Gerais) and Costa Rica (Tur- 
rialba). Koebele introduced T. echion to Hawaii from Mexico about 
1902 to control this weed (Swezey, 1913), and it was later introduced 
(1922-1923) (now extinct) to the Fiji Islands for the same reason (Rob- 
inson, 1975). Other larval foodplants of T. echion are the flowers of 
Cordia sebestena (Boraginaceae), Datura arborea (Solanaceae), Sola- 
num nodiflorum (Solanaceae), and S. sanitwongsei in Hawaii (Zim- 
merman, 1958), and Mangifera indica (Anacardiaceae) in Brasil (Lima, 


68 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


1936, 1947). Zimmerman (1958) and Lima (1947) list parasitoids of 
this species. 


Taxonomic note: The ventral wing pattern of T. echion is remarkably similar to that 
of Strymon basilides (Geyer), and has been a continuing source of confusion. We follow 
Hewitson (1862-1878), Godman & Salvin (1887-1901), Draudt (1919-1920), and Clench 
(1961) in considering T. echion to be a senior synonym of Thecla crolus Cr., and not 
a senior synonym of S. basilides (but see Kaye [1908] and Lathy [1926] for a different 
opinion). T. echion lacks the red-orange spot of S. basilides located dorsally at the 
margin of cell Cu,-Cuy. 


Larval foodplant records of T. echion and S. basilides have been 
somewhat confused as a result of the difficulty in identifying these 
species (e.g. Harris, 1927; Carter, 1933; Ehrlich & Raven, 1964). To 
date, however, T. echion has been recorded reliably only from dicots, 
and S. basilides only from monocots (see below). 

“Thecla” mathewi Hew.: A larva feeding on a fallen corolla of Cy- 
dista sp. (Bignoniaceae) pupated on 18 June 1979. A female eclosed 
before 4 July 1979. 

Cyanophrys herodotus (F.): We found a green larva on the leaves 
of Mikania sp. (Compositae) on BCI on 7 May 1980. No flowers were 
present on the plant, and the larva was successfully reared on the 
leaves. Pupation took place on 13 May; a male eclosed 25 May (Aiello 
lot 80-47). 

We observed a female bending her abdomen into the flowers of the 
introduced plant, Clerodendron paniculatum (Verbenaceae) on BCI 
on 1 August 1977 but could find no eggs. Females laid single eggs 
next to a bud of Lantana camara (Verbenaceae) on BCI on 7 June 
1979 and on a flower stalk of Cornutia grandifolia (Verbenaceae) in 
Gamboa on 18 June 1979. Neither egg was successfully reared, but 
we infer that flowers of verbenaceous plants are an important larval 
food of this common species in Panama. 

In Brasil, C. herodotus feeds on the flowers of Mangifera indica 
(Anacardiaceae) (three references in Silva et al., 1967-1968). 

“Thecla” near enenia Hew.: We found a yellow and brown larva 
on the yellow flowers of Mascagnia hippocratioides (Malpighiaceae) 
on BCI, 31 May 1980. After it molted on 2 June we gave it flowers of 
a cultivated plant, Hibiscus rosa-sinensis (Malvaceae) (flowers of 
Mascagnia were no longer available). The larva ate the Hibiscus flow- 
ers and pupated 13 June. A small male eclosed at 1400 hours on 24 
June. After eclosing the butterfly extruded and withdrew the brush 
organs (sensu Eliot, 1973) at the tip of its abdomen (Aiello lot 80-62). 


Taxonomic note: This undescribed species is closely related to enenia Hew. Males 
possess a scent pad at the end of the forewing discal cell; whereas, males of enenia 


Supplement to 
Journal of the Lepidopterists’ Society 
36(3), 1982 


Erratum for 36(2) 1982, pp. 65-75 


FOODPLANT AND OVIPOSITION RECORDS FOR 
PANAMANIAN LYCAENIDAE AND RIODINIDAE 


ROBERT K. ROBBINS AND ANNETTE AIELLO 


The printer inadvertently omitted the following paragraphs on page 
69 from the above-titled article. This erratum is printed on gum- 
backed paper which may be moistened and affixed to the bottom 
of page 69. 


“Thecla” hesperitis (B. & D.): We found two larvae inside fallen 
corollas of the liana Cydista sp. (Bignoniaceae) on BCI on 27 May 
1979. Three nematomorph worms emerged from one larva on 30 May. 
The other larva molted to the final instar on 1 June, and pupated on 
15 June. Eclosion occurred 27 June (Aiello lot 79-78). 

Beutelspacher (1972) reared this widespread, common species from 
the leaves of Tillandsia caput-medusae (Bromeliaceae) in Mexico. 

Strymon basilides (Geyer): A female laid an egg on an inflorescence 
bract of Heliconia latispatha (Musaceae) in Gamboa on 16 October 
1979. We had found literally hundreds of eggs on the flowers and 
bracts of this common species and on the bracts of H. wagneriana 
during August 1979 when these plants first blossomed. Although H. 
latispatha continued to produce flowers until at least January, we 
found only occasional eggs on this plant after mid-September. Larvae 
brought into the lab on 6 & 8 September 1979 fed on flowers of H. 
latispatha by boring into them. One pupated on 15 September and 
another on 19 September. They eclosed as females on 23 & 27 Sep- 
tember, respectively. 


VOLUME 36, NUMBER 2 69 


lack the scent pad. Otherwise, the two species are similar; future identification should 
pose no problems. We have seen specimens of enenia from Paraiba (Brasil), “Amazon, ’ 
and Guiana. The new species is known from Panama and Honduras (Museum of Com- 
parative Zoology). 


“Thecla” ericusa Hew.: Kim Steiner reared a male and female of 
this common, widespread species on flowers of Stigmaphyllon lin- 
denianum (Malpighiaceae). A larva that he found on 11 November 
1977 at Gatun pupated on 19 November and eclosed as a female. A 
larva that he found on 8 April 1980 at Frijoles pupated on 13 April 
and eclosed as a male on 21 April. T. ericusa has been reared in Brasil 
from flowers of Antigonum leptopus (Polygonaceae) (Lima, 1947), 
where it is parasitized by a wasp of the genus Tetrastichus. In addi- 
tion, there are four specimens in the U.S. National Museum from 
Trinidad and Tobago that were reared on the flowers of Dioclea gui- 
nensis and Gliricidia sp. (Leguminosae). 


Taxonomic note: Identification of species in the spurina-brescia species groups (to 
which T. ericusa belongs) has been difficult (Clench, 1961, 1970). Draudt (1919-1920) 
noted that the proximal part of the duplex male scent pad of T. ericusa “has disappeared 
except some traces of it.” This character is distinctive and is the primary basis of the 
present identification. Although the ventral wing pattern of the Brasilian type specimen 
(probably from the area around Rio de Janeiro) is considerably different from those in 
Panama, specimens with “intermediate” wing patterns occur in Bolivia, Colombia, and 
Venezuela. Male and female genitalia are indistinguishable among localities and differ 
only slightly from those of the apparently parapatric brescia Hew. 


70 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


We watched a female lay a number of eggs on Xiphidium caeruleum 
(Haemodoraceae) in Gamboa on 21 September 1979 and found three 
larvae which had bored into the immature fruit of this plant. A larva 
pupated on 8 October and eclosed as a female on 17 October. 

Larvae of S. basilides appeared to have Newcomer's glands on the 
7th abdominal segment (Newcomer, 1912), but we found no ants as- 
sociated with them. We reared no parasitoids, although others have 
(Harris, 1927; Carter, 1933; Lima, 1947). 

This species has been raised on the bromeliads Ananas cosmosus 
(=A. sativus) and Aechmea bracteata (Harris, 1927; Carter, 1933; 
Beutelspacher, 1972; Silva et al., 1967-1968, list 38 Brasilian records) 
and will probably be recorded from the flowers and fruits of other 
monocots. 


Taxonomic note: Confusion of foodplant records due to misidentifications of S. 
basilides and T. echion are noted under T. echion. A second source of confusion is that 
a few undescribed sibling species of S. basilides occur in various parts of its range, 
and some of the numerous records of pineapple (A. cosmosus) as a larval foodplant 
may refer to one or more of these sibling species. 

The morphology of the male and female genitalia of basilides as well as adult behavior 
clearly indicate a close relationship with melinus Hbn., the type species of Strymon. 
Basilides will eventually be placed in Strymon or a closely related genus. 

S. basilides has been misspelled as S. basalides by many workers (see Comstock & 
Huntington, 1959). 


Strymon yojoa (Reakirt): We observed females bending their ab- 
domens into flowers and buds of Desmodium axillare (Leguminosae) 
on BCI on 19 June and 15 July 1977 but could not find any eggs. On 
19 July 1977, however, we found three larvae boring into the fruits 
of D. axillare where we had seen oviposition behavior. Although the 
larvae apparently had Newcomer's glands, no ants were tending them. 
The larvae fed in the lab on fruits and flowers of D. axillare and 
pupated on 23-27 June. Eclosion occurred on 4 July (two males) and 
7 July (sex undetermined). 

A female deposited a single green egg on an unopened bud of Koh- 
leria tubiflora (Gesneriaceae) in Gamboa on 3 December 1979. We 
took the egg back to the lab but could not find the larva after it hatched. 
Although we found two eggshells on this plant, plus damage to buds 
and flowers typical of that done by lycaenids, we could not find any 
larvae. 

This species has been reared previously from Hibiscus tubiflorus 
and Hibiscus sp. (Malvaceae) in Mexico (Kendall, 1975). 

Michaelus vibidia (Hew.): We collected a yellow larva with brown 
stripes in a fallen corolla of Pithecoctenium crucigerum (Bignoni- 
aceae) on BCI on 6 June 1978. The larva pupated on 10 June. The 
adult eclosed 24 June (Aiello lot 78-65). 


VOLUME 36, NUMBER 2 71 


Pseudolycaena damo (Druce): A female oviposited on a stipule of 
Pterocarpus sp. (Leguminosae) on BCI on 7 June 1979. The egg 
hatched on 11 June, but the larva died two days later. This species 
has been reared from Croton niveus (Euphorbiaceae) in Mexico (Ken- 
dall, 1975), but on BCI we found no immature stages of P. damo on 
Croton bilbergianus despite extensive searching. 

“Thecla” hemon (Cramer): A female laid three eggs on twigs of 
Inga pezizifera (Leguminosae) on BCI on 8 June 1979. We found five 
other eggs on this plant. Although four eggs hatched, none was suc- 
cessfully reared. We also found three last instar larvae, presumably 
T. hemon, feeding on the leaves of this same plant on 15 June 1979, 
but they produced tachinid fly puparia from which no adult flies 
emerged. Previously recorded larval foodplants for T. hemon are Inga 
sp. in Brasil (Muller, 1878; Hoffman, 1930; Silva et al., 1967-1968) 
and the young shoots of Theobroma cacao (Sterculiaceae) in Trinidad 
(Guppy, 1904). 


Riodinidae 


Argyrogrammana crocea G. & S.: We found a flat orange-brown 
larva between two overlapping leaves of Rheedia edulis (Guttiferae) 
on BCI on 28 April 1979. The larva fed by scraping the leaf surface 
without disturbing the veins, which contain a white latex. The larva 
stopped eating and turned orange on 2 May. It made a silk girdle, 
consisting of numerous radiating strands, and pupated on 5 May. The 
pupa resembled the larva in color and pattern. Eclosion occurred on 
the evening of 17 May (Aiello lot 79-30). 

Audre domina Bat.: A female laid four blue eggs on a twig of Vismia 
baccifera (Guttiferae) near several ants (Ectatomma sp.) that were 
tending membracids on Pipeline Road (five miles north of Gamboa) 
on 11 June 1977. An ant approached the female, and she flew away. 
We found 10 other eggs on this plant but found neither eggs nor ants 
on neighboring plants of the same species. The eggs hatched on 24 
June, and the larvae died three days later. 

A female flew about plants of Turnera panamensis (Turneraceae) 
on BCI on 18 May 1979 but did not land until she encountered a 
group of Ectatomma tuberculatum. As soon as she landed, several 
ants approached her. She flew away but returned several seconds 
later. This time as the ants approached her abdomen, she turned and 
presented them with her opened wings. While the ants repeatedly 
attacked her wings with their mandibles, she laid several blue eggs 
on the branch among some older darker ones. She then flew off to 
another group of ants. Four of the older eggs hatched on 19 May and 


TZ JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


molted on 23 May. These second instar larvae, which resembled ant 
brood, did not feed and died a week later. The eggs laid on 18 May 
hatched on 31 May, but the larvae died shortly afterwards. 

Last instar larvae of two species of Audre (=Hamearis, in part) 
pupate and diapause in ant nests (Bruch, 1926; Bourquin, 1953; see 
Hemming, 1934 for generic taxonomy), and we suspect that A. do- 
mina, which is apparently single-brooded in Panama, also has an ob- 
ligate myrmecophilous relationship, including diapause in ant nests. 

Thisbe irenea Stoll: Females oviposited on leaves and twigs of Cro- 
ton bilbergianus (Euphorbiaceae) on BCI on 15 June 1977, on 6, 10, 
& 13 July 1977, and on 29 May 1979. We also found numerous larvae 
feeding on the leaves of this plant and reared three males and three 
females. Larvae fed only on younger leaves that had a pair of secreting 
extra-floral nectaries at their base on the underside of the leaf. Be- 
tween feeding bouts, larvae rested along major leaf veins with their 
heads next to a nectary. We found at least five ant species tending 
both the nectaries and the larvae, although larvae are easily reared in 
the lab without ants. The larvae have pairs of “tubercles,” which are 
occasionally everted, near their head and at their caudal end. These 
glands may be homologous with the “honey glands” and “eversible 
tubercles” reported by Ross (1966). Pupae have polyp-like glands that 
are tended by ants and have two black spots that give the appearance 
of holes through which parasitoids emerged. Pupae of the African 
lycaenid Argiolaus maesa also have spots that resemble parasitoid 
emergence holes (Hinton, 1955). Parasitoids of T. irenea include a 
chalcidid (Spilochalcis sp.) and a tachinid fly. In addition, one tach- 
inid puparium had two small holes in it indicating that a hyperpar- 
asitoid had emerged from it. 


DISCUSSION 


Most of the lycaenids we reared oviposit and feed on reproductive 
parts of plants. A notable exception is A. aetolus, which appears to 
be an obligate foliage feeder; in both the field and lab, females do 
not oviposit and larvae do not feed on flowers and fruits of their so- 
lanaceous foodplants. T. hemon and P. damo may also feed primarily 
on foliage. We also found late instar larvae of T. echion and C. her- 
odotus feeding on leaves, but females of these species have been seen 
ovipositing only on plant reproductive parts. On the other hand, we 
have never seen riodinids ovipositing on flower-buds, flowers, or fruits. 

The biological consequences of flower-feeding for lycaenid butter- 
flies, particularly in relation to polyphagy, have not been explored. 


VOLUME 36, NUMBER 2 73 


We tentatively propose three possible consequences that merit fur- 
ther investigation. First, fower-feeding may allow lycaenids, at least 
as early instar larvae, to feed on plants whose leaves are not available 
to butterfly larvae (perhaps due to chemical defenses). For example, 
Ehrlich & Raven (1964) list plant families including the Bignoniaceae, 
Gesneriaceae, and Begoniaceae that are not used, or are under-uti- 
lized by butterfly larvae. We recorded three flower-feeding lycaenid 
species (M. vibidia, T. hesperitis, and T. mathewi) on Bignoniaceae 
and one oviposition record (S. yojoa) on a flower in the Gesneriaceae. 
In addition, Zikan (1956) reared Thecla azaria Hew. from the flowers 
of a plant in the Begoniaceae. Many more records will be needed 
before we properly can assess the range of larval foodplants used by 
these insects. For evident reasons, we consider it imperative that 
foodplant records include the plant part fed upon. 

A second consequence of flower-feeding is that it may allow indi- 
vidual lycaenid species to feed on a variety of unrelated plants. The 
range of seemingly unrelated foodplants used by some of the flower- 
feeding lycaenids discussed in this paper is striking and would be 
considered highly unusual for butterflies in other groups. In addition, 
one of the most polyphagous butterfly species known (Strymon me- 
linus on 46 genera in 21 families [Howe, 1975]) is a flower-feeding 
lycaenid. Such polyphagy would be particularly advantageous to 
weedy, multivoltine species that frequently encounter changing hab- 
itats or seasons. _ 

A third possible consequence of flower-feeding is that the abun- 
dance and diversity of adult lycaenid butterflies might “track” the 
abundance and diversity of flowering plants. The peak of flowering 
on BCI occurs at the end of the dry season (February to May) (Croat, 
1978, p. 35), which corresponds to an increase of about two orders of 
magnitude in the abundance of lycaenid butterflies, at a time when 
many species in other butterfly families show a marked decrease in 
abundance (pers. obs.). A consequence of dry season abundance, in 
turn, is dispersal by strong, sustained dry season trade winds (Robbins 
& Small, 1981). 

Myrmecophily has been recorded in many lycaenid species (e.g. 
Hinton, 1951) but for only seven species of Neotropical riodinids (Cal- 
laghan, 1977). We note that in addition to the records presented in 
this paper, we have encountered myrmecophilous larvae of at least 
five other riodinid species but were unable to rear them. We agree 
with Callaghan (1977) that a large proportion of the Neotropical rio- 
dinids will prove to be myrmecophilous. 


74 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


ACKNOWLEDGMENTS 


We thank T. Antonio, R. Dressler, B. Hammel, and K. Steiner for help with plant 
identifications; G. B. Small for identifying some of the butterfly specimens; C. J. Cal- 
laghan and H. K. Clench for clarifying some taxonomic questions on these butterflies; 
L. Miller and J. Miller of the Allyn Museum for providing photographs of type speci- 
mens in the British Museum; H. Hespenheide for identifying a chalcidid; and F. S. 
Chew and R. E. Silberglied for advice on aspects of the paper. We are particularly 
grateful to Boyce A. Drummond III and Kim Steiner for allowing us to use their un- 
published foodplant records. The senior author acknowledges the support of a Smith- 
sonian Institution post-doctoral fellowship. 


LITERATURE CITED 


BEUTELSPACHER, C. R. 1972. Some observations on the Lepidoptera of bromeliads. 
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BOURQUIN, F. 1953. Notas sobre la metamorfosis de Hamearis susanae Orfila, 1953 
con oruga mirmecofila (Lep. Riodin.). Rev. Soc. Ent. Argentina 16:83-87. 

BREEDLOVE, D. E. & P. R. EHRLICH. 1968. Plant-herbivore coevolution: Lupines 
and lycaenids. Science 162:671-672. 

BRUCH, C. 1926. Orugas mirmecofilas de Hamearis epulus signatus Stich. Rev. Soc. 
Ent. Argentina 1:1-9. 

CALLAGHAN, C. J. 1977. Studies on restinga butterflies. I. Life cycle and immature 
biology of Menander felsina (Riodinidae), a myrmecophilous metalmark. J. Lepid. 
Soc. 31:173-182. 

CARTER, W. 1933. Notes on two pests of pineapple not known in Hawaii. Proc. Haw. 
Entomol. Soc. 8:395-397. 

CLENCH, H. K. 1961. Theclini. In How to Know the Butterflies. P. R. Ehrlich & A. 
H. Ehrlich. W. C. Brown Co., Dubuque, Iowa. 262 pp. 

1970. Generic notes on two hairstreaks new to the United States (Lycaenidae). 
J. Lepid. Soc. 24:56-59. 

COMSTOCK, W. C. & E. I. HUNTINGTON. 1959. An annotated list of the Lycaenidae 
(Lepidoptera, Rhopalocera) of the Western Hemisphere. J. New York Entomol. 
Soc. 67:59-96. 

CroAT, T. B. 1978. Flora of Barro Colorado Island. Stanford University Press, Stan- 
ford, CA. 943 pp. 

DOLINGER, P. M., P. R. EHRLICH, W. L. FIrcH & D. E. BREEDLOVE. 1973. Alkaloid 
and predation patterns in Colorado lupine populations. Oecologia 13:191-204. 
DOWNEY, J. C. 1962. Host-plant relations as data for butterfly classification. System- 

atic Zoology 11:150-159. 

DraupT, M. 1919-1920. Thecla F. In Macrolepidoptera of the World. vol. V. The 
American Rhopalocera. Ed. A. Seitz. Alfred Kernen Verlag, Stuttgart. 1140 pp., 
194 pl. 

EHRLICH, P. R., D. E. BREEDLOVE, P. L. BRUSSARD & M. SHARP. 1972. Weather and 
the “regulation” of subalpine populations. Ecology 53:243-247. 

EHRLICH, P. R. & P. H. RAVEN. 1964. Butterflies and plants: A study in coevolution. 
Evolution 18:586-608. 

ELIOT, J. N. 1973. The higher classification of the Lycaenidae (Lepidoptera): A ten- 
tative arrangement. Bull. Brit. Mus. (Nat. Hist.) Entomol. 28:6. 

GODMAN, F. D. & O. SALVIN. 1887-1901. Biologia Centrali-Americana. Insecta. Lep- 
idoptera-Rhopalocera. vol. II. Privately published. 782 pp. 

Guppy, J. 1904. Notes on the habits and early stages of some Trinidad butterflies. 
Trans. Entomol. Soc. Lond. 1904:225-228. 

Harris, W. V. 1927. Ona lycaenid butterfly attacking pineapples in Trimadael B.W.I. 
Bull. Ent. Res. 18:183-188. 

HEMMING, F. 1934. Notes on the types of three genera proposed by Jacob Hubner 
in the first volume of his Sammlung Exotische Schmetterlinge, with a definition 
of a new genus. Entomologist 67: 156-157. 


VOLUME 36, NUMBER 2 Ne 


HEWITSON, W. C. 1862-1878. Illustrations of Diurnal Lepidoptera. Lycaenidae. John 
van Voorst, London. 282 pp., 110 pl. 

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1947. Sobre endoparasitos de Thecla basilides (Lep., Lycaenidae). Ann. Acad. 
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ROBBINS, R. K. & G. B. SMALL, JR. 1981. Wind dispersal of Panamanian hairstreak 
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of Hawaii Press, Honolulu. 456 pp. 


Journal of the Lepidopterists’ Society 
36(2), 1982, 76-82 


DIFFERENTIAL GROWTH AND UTILIZATION OF THREE 
FOODPLANTS BY FIRST INSTAR LARVAE OF 
CITHERONIA REGALIS (SATURNIIDAE) 


C. BROOKE WORTH 
R.F.D., Delmont, New Jersey 08314 


AUSTIN P. PLATT & THOMAS F.. WILLIAMS 


Department of Biological Sciences, University of Maryland Baltimore County, 
5401 Wilkens Avenue, Catonsville, Maryland 21228 


ABSTRACT. One hundred larvae of Citheronia regalis (Saturniidae) were reared 
singly in petri dishes through the first instar on three different foodplants. They grew 
most rapidly on persimmon (Diospyros virginiana), less so on sweetgum (Liquidambar 
styraciflua), and most slowly on wing-rib sumac (Rhus copallina). Survival was high 
on persimmon and sweetgum, but on sumac it was only 67 percent, with many larvae 
refusing to feed, and others dying after ingesting some food. Frass pellet counts done 
on 12 larvae indicated that persimmon was the most advantageous plant for larval 
growth, with sweetgum being of intermediate food value. Sumac may contain repellent 
or toxic chemicals which adversely affect larval growth. The differences in foodplant 
suitability apparently are genetic in this moth strain. These results further support the 
theory that neotropical ebonies (Diospyros) and C. regalis represent a co-evolved plant- 
herbivore relationship of long-standing, with the more temperate plant genera (Liquid- 
ambar and Rhus) having been exploited as alternate food sources only in comparatively 
recent times. 


Among insects herbivory is widespread, with most species accept- 
ing a limited range of foodplants and exhibiting preferences for only 
one or two of these (Matthews & Matthews, 1978). Feeding responses 
are elicited by chemo-sensory cues (Dethier, 1966, 1970a & b; Barton- 
Browne, 1975), and a number of experiments involving lepidopterous 
larvae show clearly that such preferences can be induced and modi- 
fied by prior experiences (Jermy et al., 1968; Feeny, 1970; Hanson, 
1970, 1976; Hansen & Dethier, 1973). The degree to which plants 
and insects co-evolve has been intensively studied in several systems 
involving specific plant and lepidopteran groups (Ehrlich & Raven, 
1964, 1967; Gilbert, 1971; Gilbert & Raven, 1975). 

The polyphagous larvae of Citheronia regalis (Fabricius) (Saturni- 
idae) (royal walnut moth) recently have been shown to exhibit differ- 
ential growth and development on three genera of foodplants, rep- 
resenting three distinct plant families (Worth et al., 1979). These 
foodplants are persimmon (Diospyros virginiana L., Ebenaceae), 
sweetgum (Liquidambar styraciflua L., Hammamelidaceae), and wing- 
rib sumac (Rhus copallina L., Anacardiaceae). In that paper we pos- 
tulated that persimmon and C. regalis were most compatible because 
of a long-standing, plant-herbivore relationship of neotropical origin; 
whereas, the latter two foodplants may have been exploited later by 


VOLUME 36, NUMBER 2 WET 


this moth species as its geographic distribution extended into tem- 
perate regions. 

The present paper will explore this hypothesis further by examin- 
ing differential growth and leaf utilization among first instar C. regalis 
larvae on these same three foodplants. The demonstration of such 
feeding preferences in the young larvae at the onset of their leaf feed- 
ing will strengthen this hypothesis by indicating that such prefer- 
ences have become genetically incorporated in the moth strain we 
have been studying. The previous experiments had not been de- 
signed to test the early larval feeding responses of the royal walnut 


moth. 


MATERIALS AND METHODS 


Ten freshly eclosed C. regalis females from the Eldora (Cape May 
Co.), New Jersey stock were tethered overnight (Worth, 1980) to at- 
tract either wild, or marked and released, males during late June and 
July 1979. Thirty fertile eggs from each female were placed singly in 
petri dishes lined with moist filter paper. Ten eggs each were placed 
on small leaf cuttings of persimmon, sweetgum, and sumac. Thus, a 
total of 100 eggs (including ten from each one of the females) was 
tested on each foodplant (300 in all), thereby maximizing the genetic 
variance within each subclass of the experiment. 

Nine of the P, females had been reared on persimmon, whereas the 
tenth had fed on sumac. Among the nine persimmon females, four 
bred with released males, which also had fed on persimmon, and the 
remaining five bred with wild males, whose larval foodplants were 
unknown. The tenth female (reared on sumac) likewise bred with a 
released male, which had been reared on persimmon. Due to the 
heavy predominance of persimmon among the P, moths of both sexes, 
we have treated the broods as a fixed, rather than as a random, variable 
in the statistical analyses employed in this paper. 

The petri dishes were examined daily to determine whether or not 
the larvae were feeding and if they required additional leaf material. 
All larvae were observed until they either molted to the second instar, 
or until they died. Four larvae on each foodplant (a total of 12 in all) 
were chosen at random from among the 100 larvae, and their frass 
pellets were counted daily in an attempt to relate leaf intake and 
relative assimilation to the duration of the first instar. Due to time 
limitations, larger numbers of larvae were not used to obtain these 
data. The small size and weights of these hatchling larvae likewise 
kept us from obtaining accurate larval and fecal pellet weights. How- 
ever, since the fecal pellets are of uniform size the numbers produced 
by each larva represent a good substitute criterion. 


78 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 1. Effects of foodplant and brood on the duration of first instar (days) among 
262 Citheronia regalis larvae, with a 2-way ANOVA test and mean pair comparisons. 
The samples represent ten 1979 broods of the Eldora, NJ stock. (The sample sizes are 
given in brackets.) 


FOODPLANT 
Persimmon Sweetgum Sumac 
(PR) (SG) (SM) 
Duration of first instar (x + S.E.) 4.77 + 0.10 5.04 + 0.06 6.69 + 0.18 
[95] [100] [67] 
ANOVA of duration of first instar 
(Foodplants and Broods are fixed) df Mean square F-statistic P 
Between foodplants (F) ») 77.79 92.33 P < 0.005 
Between broods (B) 9 3.58 4.25 P < 0.005 
Interaction (F x B) 18 2.80 3.32 P < 0.005 
Within subclasses (individuals) 232 0.84 = au 


If the means (x) are ordered from largest to smallest, Duncan’s Multiple Range Tests show that the three means 
differ significantly from each other: PR < SG < SM (P < 0.05). 


Our data have been analyzed using separate 2-way analysis of vari- 
ance (ANOVA) tests on: 1) larval development time (duration of the 
first instar) and moth brood; and on 2) the number of frass pellets 
produced daily by the 12 larvae separated for this purpose. Due to an 
excessive number of zeros present in some of the daily frass count 
data, we performed a log transformation on the count data before 
analyzing them. Following both analyses, Duncan’s Multiple Range 
Tests were used to test for statistical significance between individual 
mean pairs. Our statistical procedures follow those given by Freund 
et al. (1960). 


RESULTS 


Larval survival through the first instar was high (95-100%) on both 
persimmon and sweetgum but was much lower (only 67%) on sumac 
(Table 1). On the two aforementioned plants all larvae began to feed, 
as soon as their integuments had hardened. Those larvae placed on 
sumac, however, often wandered off the leaves, as if looking for another 
food source. Although a few began to feed immediately, the majority 
of larvae exhibited this wandering behavior. Many of the wanderers 
eventually returned to the sumac leaves and began to feed, although 
sparingly. Some never fed at all and starved to death while clinging 
to the presumably edible substrate. Each of the five larvae which died 
while feeding on persimmon was from a different brood. All fed well 
at first, and the cause of their deaths was not evident. 

Table 1 shows the effects of foodplant and brood on the duration 
of first instar among the 262 surviving larvae. All three sample means 


VOLUME 36, NUMBER 2 79 


TABLE 2. Effects of foodplant and development time (days) on the number of frass 
pellets produced by 12 randomly chosen first instar Citheronia regalis larvae, with a 
2-way ANOVA test and mean pair comparisons. The frass count data have been sub- 
jected to log transformation for analysis. (The total number of pellets is in brackets.) 


FOODPLANT 
Persimmon Sweetgum Sumac 
Actual x (PR) (SG) (SM) 
aE Splde La DART ADEE Pe See ETS ee 
x No. of frass pellets produced x+S.E.of 1817 + 3.63 27.88 + 5.00 30.25 + 3.36 
over a six day period by four transformed [436] [669] [726] 
larvae on each foodplant data 7.88 + 1.01 IPG) SE ILI 26.70 + 1.01 
ANOVA of No. of frass pellets 
produced (Foodplants and 
days are fixed) df Mean square F-statistic Pp 
Between foodplants (F) 2 1.7183 39.69 P < 0.0005 
Between days (D) 5 3.16 72.54 P < 0.0005 
Interaction (F x D) 10 0.85 19.61 P < 0.0005 
Within subclasses (error) 54 0.04 — _— 


Duncan’s Multiple Range Tests show that the three log, x’s (and consequently the other means, as well) differ 
significantly from each other: PR < SG < SM (P < 0.05). 


are statistically significant from each other, with the larvae on persim- 
mon exhibiting the most rapid growth and those on sumac the slowest 
growth rate. The between brood effects are also significant in this 
analysis, as is the foodplant x brood interaction term. Thus, larvae 
from separate broods exhibit distinctive growth rates, and these rel- 
ative rates vary differentially, depending on the type of foodplant 
utilized. 

The frass pellet count results are given in Table 2. The numbers of 
droppings produced daily during the first instar by four larvae feeding 
on persimmon is significantly lower than the number produced by 
the four larvae feeding on either sweetgum, or sumac. Once again, all 
three means exhibit statistical significance and, likewise, both the 
effect between days and the foodplant x days interaction term are 
statistically significant for the reasons given above. 

The four larvae feeding on persimmon required only four days to 
complete the first instar, and they produced an average of 109 frass 
pellets. Those on sweetgum required five days in this stage and pro- 
duced an average of 167 droppings. On sumac the first instar interval 
lasted six days, and the four larvae had a mean production of 182 fecal 
pellets. 


DISCUSSION AND CONCLUSIONS 


These results confirm and extend the conclusions of our earlier 
experiments. Persimmon represents the optimal foodplant for the E]- 


80 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


dora, New Jersey stock of C. regalis when compared to either sweet- 
gum or wing-rib sumac. That sweetgum occupies an intermediate posi- 
tion is more clearly shown than in our former experiments, while 
sumac stands out as an inferior food resource. The moth larvae exhibit 
differential responses to these three foodplants immediately as they 
begin to feed. Our results further support the hypothesis that the 
persimmon-Citheronia relationship is one of long standing and one 
which has become highly adaptive. 

The frass counts suggest that the three foodplants provide different 
nutritional values, since larvae on persimmon did not have to eat as 
much to attain full size. Larvae on sweetgum had to eat half again as 
much as the former ones did to achieve the same growth. Although 
these tiny larvae were not weighed or measured, by inspection those 
on persimmon and sweetgum attained the same average size, while 
those surviving on sumac, despite feeding longer and more volumi- 
nously, were smaller when entering their first ecdysis. Such differ- 
ential utilization and assimilation of the three foodplant genera by 
freshly hatched first instar larvae strongly suggests that the New Jer- 
sey strain of C. regalis is genetically more compatible with persim- 
mon than it is with either of the two other foodplant genera. 

These findings suggest that the food substances and nutrients re- 
quired by this larval strain of C. regalis can be obtained more easily 
from persimmon than from either sweetgum or wing-rib sumac. The 
food compounds and nutrients ingested from leaves belonging to the 
later two plants are more difficult for the larvae to extract and utilize 
metabolically. Consequently, the regalis larvae are forced to expend 
more energy ingesting greater amounts of leaf material to accomplish 
the same amount of assimilation and growth when feeding on sweet- 
gum or wing-rib sumac, thus, reducing their ecological efficiencies as 
plant predators. 

Also, repellents such as toxic chemicals are present in sumac, de- 
spite the ability of some larvae to subsist on this plant. Perhaps it is 
significant that some of the other species of Rhus (R. radicans L., R. 
toxicodendron L., and R. vernix L.) possess recognized poisonous 
qualities (such as tannins and lacquer-like substances contained in 
the leaves). In spite of this, at least one neotropical species (Cither- 
onia splendens (Druce) is known to feed on certain Anacardine genera 
such as Shinus and Rhus (Vasquez, 1944), but this insect also uses 
walnut (Juglans) and wild cotton (Gossypium) in Arizona (Ferguson, 
1971). 

The sumacs (Rhus spp.) represent a temperate zone genus of an 
extensive tropical Family, consisting of about 50 genera and nearly 
400 species (Rodgers, 1920). There are 16-17 species of Rhus in North 


VOLUME 36, NUMBER 2 81 


America, nine of which are tree sized, including R. copallina. Both 
sweetgum and persimmon represent relict genera of trees, numerous 
species of which are found in the Eocene deposits of western North 
America, dating back about 55,000,000 years B.P. (Peattie, 1966). To- 
day, the genera Liquidambar and Rhus represent mainly temperate 
plant groupings; whereas, Diospyros is more closely affiliated with 
tropical ebonies. This, we believe, is why persimmon represents the 
better foodplant for C. regalis, a temperate moth species having nu- 
merous neotropical relatives. We suspect that C. regalis has exploited 
the temperate plant genera more recently and that Diospyros repre- 
sents the major foodplant of long-standing. 

Larvae obtained from the moth that had been reared on sumac (and 
had mated with a male reared on persimmon) did not display a higher 
survival rate on sumac than offspring of moths in which both parents 
had been reared on persimmon. Whether better adaptation to sumac 
would take place over successive generations when both parents are 
reared on sumac is a question to which the senior author will next 
direct his curiosity. 


ACKNOWLEDGMENTS 


We thank Dr. F. E. Hanson of UMBC and an anonymous reviewer for valuable 
comments on the manuscript, and Mr. P. O. Hubbell of Tuscon, Arizona for his critical 
comments relating to the evolution of the plant families discussed in the paper. Support 
for the data analyses was provided by the UMBC Computer Center. 


LITERATURE CITED 


BARTON-BROWNE, L. 1975. Regulatory mechanisms in insect feeding. Adv. Insect 
Physiol. 11:1-116. 

DETHIER, V. G. 1966. Feeding behavior. Roy. Entomol. Soc. London Symposium 3, 
pp. 46-58. 

1970a. Some general considerations of insects’ responses to the chemicals in 

food plants. In Control of Insect Behavior by Natural Products, D. L. Wood, R. M. 

Silverstein, and M. Nakajima, eds. Academic Press, NY, pp. 21-28. 

1970b. Chemical interactions between plants and insects. In Chemical Ecol- 
ogy, E. Sondheimer and J. B. Simons, eds. Academic Press, NY, pp. 83-102. 

EHRLICH, P. R. & P. H. RAVEN. 1964. Butterflies and plants: a study in coevolution. 
Evolution 18:586—608. 

1967. Butterflies and plants. Sci. Amer. 216:104-113. 

FEENY, P. P. 1970. Seasonal changes in oak leaf tannins and nutrients as a cause of 
spring feeding by winter moth caterpillars. Ecology 51:565-58 1. 

FERGUSON, D. L. 1971. In Dominick, R. B. et al., The Moths of American North of 
Mexico, Fasc. 20.2A, Bombycoidea (Saturniidae), E. W. Classey, Ltd., Middlesex, 
England. 153 pp. 

FREUND, J. E., P. E. LIVERMORE & I. MILLER. 1960. Manual of Experimental Statis- 
tics. Prentice-Hall, Inc., Englewood Cliffs, NJ. 132 pp. 

GILBERT, L. E. 1971. Butterfly-plant coevolutions: has Passiflora adenopoda won the 
selectional race with heliconiine butterflies? Science 172:585-586. 

GILBERT, L. E. & P. H. RAVEN, Eds. 1975. Coevolution of Animals and Plants. Univ. 
Texas Press, Austin, TX. 246 pp. 


82 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


HANSON, F. E. 1970. Sensory responses of phytophagous lepidoptera to chemical and 
tactile stimuli. In Control of Insect Behavior by Natural Products, D. L. Wood, R. 
M. Silverstein, and M. Nakajima, eds. Academic Press, NY, pp. 81-91. 

1976. Comparative studies on induction of food choice preferences in lepi- 
dopterous larvae. Symp. Biol. Hung. 16:71-77. 

HANSON, F. E. & V. G. DETHIER. 1973. Role of gustation and olfaction in food plant 
discrimination in the tobacco hormworm, Manduca sexta. J. Insect Physiol. 
14; 1019-1034. 

JeRMY, T., F. E. HANSON & V. G. DETHIER. 1968. Induction of specific food pref- 
erence in lepidopterous larvae. Entomol. Exp. & Appl. 11:211-230. 

MATTHEWS, R. W. & J. R. MATTHEWS. 1978. Insect Behavior. John Wiley & Sons, 
NY. 507 pp. 

PEATTIE, D.C. 1966. A Natural History of Trees of Eastern and Central North Amer- 
ica, 2nd Ed. Houghton Mifflin Co., Boston, MA. 606 pp. 

RODGERS, J. E. 1920. The Tree Book. The Nature Library. Doubleday, Page & Co., 
Garden City, NY. 589 pp. 

VASQUEZ, L. 1944. Citheronia splendens queretana subsp. nov. Anales del Instituto 
de Biologia Mexico 15:235-236. 

WorTH, C. B. 1980. An elegant harness for tethering large moths. J. Lepid. Soc. 
34:61-63. 

WoRTH, C. B., T. F. WILLIAMS, A. P. PLATT & B. P. BRADLEY. 1979. Differential 
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plants. J. Lepid. Soc. 33: 162-166. 


Journal of the Lepidopterists’ Society 
36(2), 1982, 83-86 


DYSSTROMA HERSILIATA FORM “MIRANDATA’—A 
RECESSIVE COLOR FORM (GEOMETRIDAE) 


KB BOLLE 
Canadian Forestry Service, Environment Canada, Ottawa K1A 0C3 


ABSTRACT. Two successive generations of Dysstroma hersiliata hersiliata (Gn.) 
and Dysstroma hersiliata form “mirandata’ (Tayl.) have been successfully reared in 
the laboratory. The results of selective matings have shown that D. hersiliata form 
“mirandata’” is homozygous for a recessive color form of D. hersiliata hersiliata. Rear- 
ing and overwintering techniques are described. 


Dysstroma hersiliata hersiliata was originally described by Gue- 
née in 1857 and D. h. form “mirandata” by Taylor in 1910. Distinct 
differences between the two can be seen in the color of the median 
and antemedian bands. Specimens of D. hersiliata have a dark grey 
median band and a deep orange-yellow antemedian band with paler 
inner and outer margins; those of D. h. form “‘mirandata” have a yel- 
low-brown median band with dark grey costal edge and an anteme- 
dian band which is a slightly darker orange-yellow than that of her- 
siliata. Dysstroma h. form “mirandata” is similar to D. h. cervinifascia 
(WI1k.) except for the antemedian band which in cervinifascia appears 
as a dark greyish yellow-brown band with a yellow-brown median 
Margin. 

Dysstroma hersiliata cervinifascia and D. h. form “mirandata” seem 
to have the same geographic distribution as D. hersiliata, in fact a 
collection made at Mistassini, Quebec by J. R. McGillis on 8 August 
1956, have turned up hersiliata, h. cervinifascia and h. form “mi- 
randata’ specimens. The male and female genitalia of the above 3 all 
appear to be similar, which indicates a strong possibility that h. cer- 
vinifascia may be just another color form of hersiliata as well. Future 
rearings may prove or disprove this. 


MATERIALS AND METHODS 


Two adult females, one D. hersiliata hersiliata and one D. hersi- 
liata form “mirandata” were captured 5 July 1978 at Dunrobin Ontario 
by Dr. Eugene Munroe. These were placed in individual 32 oz waxed 
cardboard containers in which gooseberry foliage and a vial of sugar 
water with a cotton wick were placed. After the females had died, all 
the eggs were placed on a piece of #40 nylon mesh over damp ver- 
miculite in 8 oz clear plastic containers. The eggs were stored out- 
doors in a screened, shaded building until 10 October 1978, when 
the containers were placed in plastic bags and buried approximately 
22.5 cm (9 inches) underground among gooseberry plants and left to 


84 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


overwinter. On 19 April 1979, as the buds of the gooseberry were 
beginning to open, the eggs were removed from the ground, brought 
in to room temperature and placed in 8 oz plastic containers lined 
with damp tissue paper. After hatching, the larvae were placed into 
groups of 10 in 8 oz plastic containers with gooseberry foliage. Each 
resulting pupa was placed in individual 2 oz plastic vials with a small 
piece of damp wick. Soon after the moths emerged and their wings 
had dried they were placed in pairs in 32 oz waxed cardboard con- 
tainers and set up in the same manner as the original parental rearing. 

Twelve pairs were placed together for mating. The resulting eggs, 
larvae and pupae were prepared and cared for in the same manner as 
previously described. 

One of the most difficult aspects of rearing this Dysstroma species 
was overwintering the eggs successfully. Several previous attempts 
to overwinter eggs for varying lengths of time in a refrigerated envi- 
ronment had failed. However, the eggs were successfully overwin- 
tered when buried in the ground. In the Ottawa area the ground in 
the winter of 1978-79 had a good snow cover for most of the winter; 
whereas, the ground in the winter of 1979-80 was bare for most of 
the season, resulting in deep frost penetration. This may have influ- 
enced the percentage of eggs which hatched successfully in the two 
winters; approximately 44% in 1978-79 and approximately 20% in 
1979-80. Water did seep into the containers while they were buried 
in the ground both years, despite precausions taken against this; 
nevertheless, the eggs still hatched successfully. The humidity in the 
refrigerated environment was probably much too low, causing the 
previous failures. 


RESULTS 


By 10 July 1978 the original wild female D. hersiliata (79-1) had 
laid 40 eggs and the female D. h. form “mirandata” (79-2) 75 eggs. 
The eggs were laid singly, loosely attached to the foliage and bottom 
of the container. By 25 April 1979, 290 days later, 31 larvae of 79-1 
and 58 larvae of 79-2 had hatched. By 4 June 1979, 36 days later, all 
adults of the F1 generation had emerged. The 79-1 stock produced 
13 male and 10 female hersiliata offspring, (100% hersiliata), while 
79-2 produced 11 male and 13 female hersiliata, as well as 10 male 
and 18 female “mirandata”’ offspring, (46% hersiliata and 54% “‘mir- 
andata’’). Of the 12 F1 generation matings, 7 were successful in pro- 
ducing F2 offspring. These are listed in Table 1. 

There were only 2 distinct color forms from these crosses with no 
intermediates. 


VOLUME 36, NUMBER 2 


85 


TABLE 1. Results of selective matings between Dysstroma hersiliata and D. hersi- 


liata form “‘mirandata.” 


Parent Fl 


Offspring F2 


Num Number % 
ber Male Female of eggs hersiliata “mirandata”’ hersiliata 
1 hersiliata hersiliata 30 5 female 100% 
79-1 79-2 5 male 

2 hersiliata hersiliata 19 4 female 100% 
79-2 79-1 2 male 

3 hersiliata “mirandata”’ 176 15 female 9 female 50% 
79-1 79-2 19 male 25 male 

4 hersiliata hersiliata 25 1 female 50% 
79-2 79-2 1 male 

5 *’mirandata”’ “mirandata”’ 45 13 female 0% 
79-2 79-2 11 male 

6 hersiliata hersiliata 23 2 female 100% 
79-2 79-2 2 male 

ff hersiliata hersiliata 4] 100% 
79-2 79-2 2 male 

CONCLUSION 


The wild female D. h. form ““mirandata,” 79-2, produced both her- 
siliata and h. form “mirandata’” offspring demonstrating that the two 
are color forms of the same species. If one assumes that “mirandata”’ 
is the recessive color form, and judging by the 46% hersiliata and 
54% “mirandata’ F1 generation 79-2 produced, then one of the part- 
ners of 79-2 would have been homozygous for the recessive color 
characters, aa, while the other partner would have been heterozygous, 
Aa (aa X Aa). The wild female hersiliata, 79-1, produced only her- 
siliata offspring. If one assumes that hersiliata is the dominant color 
form then the partners of 79-1 could have been only two possible 
combinations, judging by the 100% hersiliata F1 generation that was 
produced. Either both partners were homozygous for dominant color 
characters, AA and AA, or one partner was homozygous, AA, and the 
other heterozygous, Aa (AA x AA or Aa X AA). In pair number 3 of 
the F1 matings a male hersiliata offspring from 79-1 was mated with 
a female “mirandata” offspring from 79-2. Since 50% of the F2 gen- 
eration was hersiliata and 50% “‘mirandata,” the F1 generation of 79-1 
probably resulted from an Aa X aa mating, producing 50% of the F1 
generation of 79-1 heterozygous and 50% homozygous for the reces- 
sive color form (Aa or aa). If the male of 79-1 had been homozygous 
for the dominant color character then all the F2 generation of the 
number 3 F1 mating would have been as the hersiliata color form. 


86 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


In pair number 5 of the F1 matings a male and female “mirandata”’ 
of 79-2 were mated. All of the F2 generations were the “mirandata”’ 
color form as would be expected if “mirandata”’ is the recessive form. 
I therefore conclude that typical D. hersiliata is a dominant color form 
and that D. h. form “mirandata”’ a recessive color form. 


LITERATURE CITED 


BARNES, W. & J. MCDUNNOUGH. 1912. Notes on Taylor’s types of Geometridae. Can. 
Entomol. 44:274. 

DyaR, H. G. 1904. The Lepidoptera of the Kootenai District of British Columbia. 
Proc. U.S. Nat. Mus. 27:136. 

FORBES, W. T. 1948. Lepidoptera of New York and Neighboring States, 2. Mem. 
Comell Univ. Agric. Exp. Stn., No. 274:136. 

ITAMIES, J. 1971. Variations in some geometrid moths (Lep. Geometridae) in Rauma 
in the years 1967-1970. Ann. Entomol. Fenn 37(4): 195-202. 

KELLY, K. L. 1965. ISCC-NBS Color name charts illustrated with centroid colors. 
Supplement to Natl. Bur. Standards Circ. 553. Washington, D.C. 

McCDUNNOUGH, J. 1927. The Lepidoptera of the Seton Lake region, British Columbia. 
Can. Entomol. 59:240. 

1939. New North American Geometridae with notes, 1 (Lepidoptera). Can. 

Entomol. 71:187-192. 

1943. Notes and descriptions of North American Geometridae. Can. Entomol. 
152 1NEPNG. 

Morris, R. F. 1980. Butterflies and moths of Newfoundland and Labrador. The Ma- 
crolepidopter. Res. Br. Agric. Can. Pub. 1691. 

MUNTZING, A. 1961. Genetics: Basic and applied. Stockholm, Sweden. 

SWETT, L. W. 1917. The genus Dysstroma Hubner. Can. Entomol. 49:64—72. 


Journal of the Lepidopterists’ Society 
36(2), 1982, 87-111 


DATES OF SELECTED LEPIDOPTERA LITERATURE FOR 
THE WESTERN HEMISPHERE FAUNA 


JOHN B. HEPPNER 


Department of Entomology, Smithsonian Institution, 
Washington, D.C. 20560 


ABSTRACT. Lepidoptera literature of 58 authors, involving Western Hemisphere 
species which have had various dates applied to specific titles, are listed and correct 
dates noted. In some cases works are given dates by sections of pages and plates. 


This selection of literature references on Western Hemisphere Lep- 
idoptera, primarily original descriptions of taxa, does not include all 
works on Lepidoptera of this fauna. Included are only those works 
that have repeatedly been dated incorrectly by subsequent authors or 
had various kinds of confusion surrounding their correct dates. In 
many cases dates of specific pages or plates are noted. 

This list was originally prepared in conjunction with the work of 
the collaborators now engaged in cataloging and reviewing the Neo- 
tropical Lepidoptera for the series, Atlas of Neotropical Lepidoptera. 
Inasmuch as various dates have often been applied to various works 
on Neotropical Lepidoptera, this list was prepared to determine the 
correct dates of certain works as best known at this time, thus ensuring 
consistency regarding dates used in the Atlas. It is hoped that this 
listing will also be of help to other authors when consulting these 
works. 

The literature is listed alphabetically and chronologically by author. 
For each of the 58 authors in the main listing, their full known name 
and dates of birth and death are provided when known. Likewise, the 
full titles to each work are given. A second section to this listing 
covers works on dates of the main citations and other works on Lep- 
idoptera. 

A number of persons helped to ensure the accuracy of dates in this 
listing, the following having been especially involved: D. S. Fletcher, 
and K. Sattler [British Museum (Natural History), London]; J. G. 
Franclemont (Cornell University, Ithaca, New York); A. Diakonoff 
(Rijksmuseum van Natuurlijke Historie, Leiden, Netherlands); G. 
Lamas (Museo de Historia Natural, Lima, Peru); and J. F. G. Clarke 
and E. L. Todd (Smithsonian Institution, Washington, D.C.). 


Selected Lepidoptera Literature 
Agassiz, Jean Louis Rodolphe (1807-1873) 


1842-1847. Nomenclator zoologicus, continens nomina systematica generum ani- 


88 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


malium tam viventium quam fossilium. Soloduri: Jent and Grassman. 12 fascicles. 
[Lepidoptera: fascicle 9-10, 1846. Index universalis, 1847. (see Bowley & Smith, 
1968; Fletcher, 1979)]. 


Berthold, Arnold Adolph (1803-1861) 


1827. Natiirliche Familien des Thierreichs. Aus dem franzosischen, mit Anmerkun- 
gen und Zusatzen. Weimar. 604 pages. [predates Latreille, 1829, of which it is a 
German translation with additions by Berthold.] 


Boisduval, Jean Baptiste Alphonse Déchauffour de (1801-1879) 


1832. Faune entomologique de l’ocean Pacifique: Lépidopteres. In, Voyage de ... 
l Astrolabe ... pendant... 1826-29, sons le commandement de M. J. Dumont d’Ur- 
ville, & c. Paris: Tastu. 267 pages, plates 1-5. [April 1832]. 


1836. Histoire naturelle des insectes: Spécies général des lepidopteres. Paris: Roret. 
690 pages, 24 plates. [type-species designations do not conform to Article 69(a) (iii) 
of the ICZN] [11 April 1836; see Cowan, 1969]. 


[1875]. Sphingides, Sesiidae, Castnides. In J. B. A. D. de Boisduval and A. Guenée, 
Histoire naturelle des insectes: Spécies général des lépidopteres hétérocéres, 1:568 
pages, 11 plates. Paris: Roret. [not 1874]. 

Busck, August (1870-1944) 
[1934]. Microlepidoptera of Cuba. Entomologica Americana 13:151—202, plates 30-36. 
[not 1933]. 

Butler, Arthur Gardiner (1844—1925) 


1869-1874. Lepidoptera exotica, or descriptions and I]lustrations of Exotic Lepidop- 
tera. London: E. W. Jansen. 190 pages, 64 plates. [Pp. 129-136 missing through 
typographical error]. 


Jun 1869 Pp. 1-8 Pls. 1-3 

Sep 1869-71 9-58 4-22 
Apr 1871 59-66 23-25 
Jul 1871 67-74 26-28 
Oct 1871 75-84 29-32 
Jan 1872 85-94 33-35 
Apr 1872 95-104 36-38 
Jul 1872 105-114 39-41 
Oct 1872 115-120 42-44 
Jan 1873 121-128 45-48 
Apr 1873 137-144 49-51 
Jul 1873 145-152 52-54 
Oct 1873 153-162 55-57 
Jan 1874 163-174 58-60 
Apr 1874 175-190 61-64 


Clereck, Carl Alexander (1710-1765) 


1759-[1764]. Icones insectorum rariorum cum nominibus eorum trivialibus, locisque 
e C. Linnaei ... Systema Naturae allegatis. Holmiae [=Stockholm]. 21 pages, 55 
plates. [1759: leties 1-12. [1764]: plates 13-55. (see Higgins, 1970)]. 


Costa, Orenzio-Gabriele (1787-1867) 


[1836]-1850. Lepidotteri. In Fauna del Regno di Napoli ossia enumerazione di tutti 
gli animali che abitano le diverse regioni di questo regno e le acque che le bagnano 
contenente la descrizione de nuovi 0 poco esattamente conosciuti. Naples: Torchi. 
446 pages, 38 plates. [dates by Direction 59 (see Tremewan, 1977)]. 


VOLUME 36, NUMBER 2 


[1836] 
1848 
1849 
1850 
1849 


Cramer, Pieter (1721-1776) 


Pp. 1-314 
315-370 
371-402 
403-418 
419-422 


89 


1775-1782. De Uitlandsche Kapellen voorkomende in de drie waereld-deelen Asia, 
Africa en America. Amsterdam: Baalde. 4 volumes, 400 plates. [Opinion 516, plus 
Cramer as author of vol. 4 by posthumous publication (see Roepke, 1956)]. 


1775 
1776 
LUPATE FE 
£79 
1780 
1780 
1781 
1782 
1782 


Curtis, John (1791-1862) 


1823-1839. British entomology; being illustrations and descriptions of the genera of 
insects found in Great Britain and Ireland: containing coloured figures from nature 
of the most rare and beautiful species, and in many instances of the plants upon 
which they are found. London. 16 vols., 770 pls. [See Sherborn & Durrant (1911): 
title page dated 1823 but plates are dated from 1824-1839. Second edition is dated 


1829-1840 from the plates. Third edition is a reprint dated 1862.] 


Vole le ppe. 132 


133-156 


Vol. 2: pp. 1-152 
Vol. 3: pp. 1-104 


105-176 


Vol. 4: pp. 1-90 


Supplement (by C. Stoll) 


91-164 
165-252 


Pp. 1-20 


Pls. 


1-84 
85-96 
97-192 

193-252 
253-288 
289-336 
337-372 
373-396 


397-400 


[Denis, Johann Nepomuk Cosmas Michael (1729-1800), and Ignaz Schiffermuller 


(1727-1809)] 


1775. Ankundung eines systematisches Werkes von den Schmetterlingen der Wie- 
nergegend. Vienna: Bemardi. 322 pp., 2 pls. [Authorship and date by Opinion 516 


(ICZN); 1776 edition with altered title page; see Sattler, 1970] 


Diakonoff, Alexey Nikolaevich 


[1968]. Microlepidoptera of the Philippine Islands. Bull. U.S. Natl. Mus. 257:1-484. 
[31 Jan. 1968, not 1967; see Clarke, 1980]. 


Donovan, Edward (1768-1837) 


1792-1813. The natural history of British insects; explaining them in their several 


states .. 


microscope. London. 16 vols., 576 pls. 


1792 
1793 
1794 
1795 
1796 
1797 
1798 
799 
1800 
1801 
1804 


Vol. 


ao 


FPOoOOoOoOnNndourhwWNrH 


Pls. 


. with the history of such minute insects as require investigation by the 


1-36 
37-72 
73-108 

109-144 
145-180 
181-216 
217-252 
253-288 
289-324 
325-360 
361-396 


90 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


1807 12 107 pp. 397-432 
1808 13 78 pp. 433-468 
1810 14 Opp: |. 469-504 
1811 15 88 pp. 505-540 
1813 16 101 pp. 541-576 


1822-1827. Naturalist’s repository, or miscellany of exotic natural history exhibiting 
... specimens of foreign birds, insects, shells, & c. London. 5 vols., 180 pls. [See 
Reynell (1917): each number published as 3 plates and accompanying text each 
month. ] 


[Apr 1822—Mar 1823] Vol. 1: viii & 198 pp. Pis.3, #36 
[Apr 1823-—Mar 1824] Vol. 2: iv & 236 pp. 37-72 
[Apr 1824—Mar 1825] Vol. 3: iv & 104 pp. 73-108 
[Apr 1825-—Mar 1826] Vol. 4: iv & 113 pp. 109-144 
[Apr 1826—Mar 1827] Vol.5: iv & 98 pp. 145-180 


Doubleday, Edward (1811-1849) 


1846-1852. The genera of diurnal Lepidoptera: comprising their generic characters, 
a notice of the habits and transformations, and a catalogue of the species of each 
genus. London: Longman. 534 pp., 67 pls. [see Hemming (1941): J. O. Westwood 
authored volumes from 1850-1852. ] 


1B Webi) Bs eS Pls. A-4 

1847 19-132 5-25 & 28 

[1848] 133-142; 145-200 26-27. 29: 31-44 

[1849] 201-242 30; 45-52; 56-58; 60-62; 64 
[1850] —[Vol. 2] 243-326 53-55; 63; 65-66 

[1851] 327-466 59: 67 

[1852] 467-534 at 


Druce, Herbert (1846-1913) 


1881-1900. Lepidoptera-Heterocera, in F. D. Godman and O. Salvin, Biologia Cen- 
trali-Americana; or, contributions to the knowledge of the fauna and flora of Mexico 
and Central America. Zoology: Insecta. London: Taylor & Francis. 3 vols. 


1881 Vol.1: pp. 1-24 

1883 25-32 

1884 S352 

1885 113-160 

1886 161-200 

1887 201-256 

1889 257-344 

1890 345-440 

1891 441-490 Pls. 1-40 [undated] 
1891 Vol. 2: pp. 1-24 

1892 25-128 

1893 129-184 

1895 185-272 

1896 273-336 

1897 337-440 

1898 441-536 

1899 537-592 

1900 593-622 Pls. 41-101 [undated] 


Drury, Dru (1725-1803) 


1770-1782. Illustrations of natural history; wherein are exhibited . . . figures of exotic 
insects, according to their different genera, & c. London: White. 3 vols. [See Opinion 
474 (ICZN)]. 


VOLUME 36, NUMBER 2 91 


1770 Vol. 1: 130 pp., 50 pls. [no scientific names] 
1773 (index to vol. 1) [plus scientific names] 
LiaS Vol. 2: 90 pp., 50 pls. 
1782 Vol. 3: 76 pp., 50 pls. 


Duponchel, Philogene Auguste Joseph (1774-1846) 


1844-1846. Catalogue méthodique des lépidopteres d'Europe pour servir de com- 
plement et de rectification a histoire naturelle des lépidopteres de France. Paris: 
Meéquignon-Marvis 523 pp. 


1844 Pp. 1-64 
1845 65-296 
1846 297-523 


Dyar, Harrison Gray (1866-1929) 
[1903]. A list of North American Lepidoptera and key to the literature of this order 
of insects. Bull. U.S. Natl. Mus. 52: 1-723. [not 1902; see Clarke, 1950]. 

Esper, Eugen Johann Christoph (1742-1810) 


1776-[1830]. Die Schmetterlinge in Abbildungen nach der Natur mit Beschreibun- 
gen. Erlangen: W. Walthers. 5 vols. [See Heppner, 1981]. 


Theil I: Die Tagschmetterlinge. 


1776 Boe l—72 Pls, 3-12 
aha 73-176 13-36 
1778 177-216 37-48 
1779 217-388 49-50 
Fortsetzung. 
1780 1-36 51-56 
1781 37-124 57-74 
1782 125-140 75-80 
1783 141-172 81-88 
1784 173-184 89-92 
1786 185-190 93 
Supplement. Theil I: Abschnitt 1. 
1789 1-60 94-101 
1794 61-68 102-106 
[1798] 69-88 107-109 
1800 89-104 110-112 
[1803-04] 105-120 113-116 
Supplement. Theil 2. 
1805 1-24 117-122 
[1805-30] 25-48 123-126 
Theil II: Die Abendschmetterlinge. 
1778 = Pls. 1-6 
1779 1-80 7-18 
1780 81-196 19-[25] 
Fortsetzung. 
1782 NOD, 26-31 
1783 213-228 32-35 
1786 229-234 36 
Supplement. Abschnitt 2. 
1789 14 37 
[1789] 5-12 [38-40] 
1794 13-16 41 
1798 17-20 — 
1800 21-40 42-46 
[1803-04] AN=52 47 


92 


Theil III: Die Nachtschmetterlinge. 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Pls. 


1-12 
13-23 & 6A 
24-31 | 
32-55 
56-79 


(Noct. 
(Noct. 
(Noct. 
(Noct. 
(Noct. 


(rev. pls. 99B & 105B) 


156-177 


(Noct. 


(rev. pls. 117a & 125b-—c) 


178-183 


184-186 
187-191 
[192-193] 
194-196 
197-198 


(Noct. 


1-12) 
13-42) 
43-60) 
61-66) 
67-76) 


77-98) 


99-104) 


Noct. 105-107) 


( 
( 
(Noct. 113-114) 
( 
( 


. 108-112) 


_ 115-117) 
. 118-119) 


Theil V: Die Spannerphalenen. [pp. 129-148 lacking by error in numbering] 


1782 Bo 136 
1783 57-104 
1784 105-136 
1785 137-272 
1786 273-396 
Supplement. Abschnitt 3. 
1789 1-36 
[1789] — 
1794 37-64 
[1798] 65-72 
1800 73-94 
[1800-07] 95-100 
[1807] 101-104 
Theil IV: Die Eulenphalenen. 
Band I. 
1786 Bo, Ieee 
1787 33-112 
1788 113-160 
1789 161-176 
1790 177-208 
1791 209-352 
1794 353-372 
Band IT. Abschnitt 1. 
1796 373-420 
1798 421-464 
[1799-1803] 465-496 
1804 497-632 
1805 633-698 
Band II. Abschnitt 2. 
1798 1-44 
[1799-1803] 45-54 
1804 55-62 
1805 63-85 
1795 se LAO 
[1795-1801] 41-196 
1801 197-220 
1803 221-244 
[1806] 245-276 


PIse 


14 


1784-1801. Die auslandischen oder die ausserhalb Europa zur Zeit in den tbrigen 
Welttheilen vorgefundenen Schmetterlinge in Abbildungen nach der Natur mit Be- 
schreibungen. Erlangen: W. Walthers. 254 pp., 63 pls. [See Hayward, 1953; Poche, 
1938]. 


1784 Pp. 1-20 Pls. 14 

1785 21-36 5-8 

1786 37-52 9-12 
1788 53-64 13-16 
1790 65-80 17-20 
7 Oil 81-96 21-24 
1792 97-144 25-36 
1793 145-160 37-40 
1796 161-192 41-46 


VOLUME 36, NUMBER 2 93 


LASIE 193-204 47-50 
1799 205-220 51-54 
1801 221-254 55-59 & 60A-D 


Eversmann, Eduard Friedrich von (1794—1860) 


1841. Fauna entomologica, quam per viginti fere annes in provinciis Volgam fluvium 
inter et montes Uralenses observavit et descriptionibus illustravit. Vol. 1: Lepidop- 
tera. Casani. 166 pp. 


1844. Fauna lepidopterologica Volga-Uralensis exhibens lepidopterorum species. 
Casani. 633 pp. [A revised edition of 1841, with added pages.] 


Fabricius, Johann Christian (1745-1808) 


1777. Genera insectorum eorumque characteres naturales ... adjecta mantissa spe- 
cierum nuper detectarum. Chilonii [=Cologne]: Bartsh. 324 pp. 


1781-[1782]. Species insectorum exhibentes eorum differentias specificas, synonyma 
auctorum, loca natalia, metamorphosin adiectis observationibus, descriptionibus. 
Hamburg & Cologne: Bohn. 2 vols. 


1781 Band 1: pp. 1-552 
Band 2: pp. 1-494 
[1782] Appendix: pp. 495-514; index pp. 515-517. 


1807. In J. C. Illiger, Die neueste Gattungs-Eintheilung der Schmetterlinge aus den 
Linneischen Gattungen Papilio und Sphinx. Mag. Insektenkunde 6:277-295. [See 
Opinion 232 (ICZN) for priority over Fabricius, 1807b.] 


1807b. Index alphabeticus in systema antliatorum genera et species continens ... 
1806. Brunsvici [=Braunschweig]: Reichard. 32 pp. 


Feisthamel, Joachim Francois Philiberto de (1791-1851) 


1839. Supplement a la zoologie du voyage de la Favorite comprenant la description 
de lépidopteres nouveaux. Mag. Zool. 9:17—26; 10 pls. [Reprinted in 1840: 13 pp., 
10 pls. Paris: Bertrand.] 


Felder, Cajetan (1814-1894), Rudolf Felder (1842-1871), and Alois Friedrich 
Rogenhofer (1831-1897) 
1865-1875. Reise der Osterreichischen Fregatte Novara um die Erde in den Jahren 
1857, 1858, 1859 unter den Behilfen des Commodore B. von Willerstorf-Urbair. 
Zoologischer Theil. Zweiter Band: Zweiter Abtheilung. Vienna. 5 parts, 140 pls. [See 
Dalla Torre, 1913; Fletcher, 1979; Higgins, 1963]. 


[1865] Heft 1 (Felder & Felder) Pp. 1-136 Pls. 1-21 
1865 Heft 2 (Felder & Felder) 137-378 22-47 
1867 Heft 3 (Felder & Felder) 379-535 48-74 
1874 Heft 4 (R. Felder) 1-10 75-107 
1875 Heft 5 (R. Felder & Rogenhofer) 1-20 108-140 


Fischer von Roslerstamm, Josef Emanuel (1787-1866) 


1834-1843. Abbildungen zur Berichtigung und Erganzung der Schmetterlingskunde, 
besonders der Microlepidopterologie als Supplement zu Treitschke’s und Hubner’s 
europaeischen Schmetterlinge, mit erlauterndem Text. Leipzig. 308 pp., 100 pls. [20 
parts]. [See Fletcher & Griffin, 1943.] 


1834 Pp. 1-16 Pls. 1-10 
1835 17-36 11-20 
1836 37-60 21-30 
1837 61-102 31-40 


1838 103-132 41-50 


94 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


1839 133-188 51-65 
1840 189-216 66-75 
1841 217-268 76-90 
1842 269-288 ; 91-95 
1843 289-308 96-100 


Freyer, C. F. (?) 
1831-1858. Neuere Beitrage zur Schmetterlingskunde mit Abbildungen nach der 
Natur. Augsburg. 7 vols., 700 pls. 


1831 Vol. 1: pp. 1-80 

1832 81-162 

1833 163-182 Pls. 1-96 [undated] 
1833 Well. 2a. 120 

1834 4\-82 

1835 83-114 

1836 115-162 Pls. 97-192 [undated] 
1836 Vols Sea. 1-80 

1837 21-52 

1838 53-124 

1839 125-166 Pls. 193-288 [undated] 
1839 Voli 4p l=20 

1840 21-82 

1841 83-94 

1842 95-166 Pls. 289-384 [undated] 
1842 Vol. 5: pp. 1-40 

1844 41-92 

1845 93-164 Pls. 385-480 [undated] 
1846 Voll6.pp 1220 

1848 21-82 

1849 83-104 

1850 105-146 

1851 147-198 

1852 (title page) Pls. 481-600 [undated] 
1858 Vol.7: pp. 1-180 Pls. 601-700 [undated] 


[no wrapper dates available for parts of volume 7] 


Godart, Jean Baptiste (1775-1825) 


1819-[1824]. Papillons. In Histoire naturelle; entomologie, ou histoire naturelle des 
crustacés, des arachnides, et des insectes, in Encyclopédie méthodique. Vol. 9. Paris. 
[See Cowan (1967, 1968) for parts credited to Latreille]. 


1819 Pp. 1-328 
[1824] 329-828 


Godman, Frederick Ducane (1834-1919), and Osbert Salvin (1835-1898) 


1879-1901. Lepidoptera—Rhopalocera. In Biologia Centrali-Americana . . . . Zoology: 
Insecta. London: Taylor & Francis. 3 vols. 


1879 Vole pppoe 56 

1880 57-88 

1881 89-168 
1882 169-224 
1883 225-288 
1884 289-360 
1885 361-400 
1886 401-487 
1887 Wolk Ze joo, I 


1889 113-184 


VOLUME 36, NUMBER 2 95 


1890 185-240 
1893 241-328 
1894 329-384 
1895 385-416 
1896 417-440 
1897 441-448 
1899 449-460 
1900 461-588 
1901 589-782 
[no date] Vole 3: ul sols: 


Gray, George Robert (1808-1872) 


1853. Catalogue of lepidopterous insects in the ... British Museum. Pt. 1: Papilion- 
idae. London. 84 pp., 15 pls. (8 Jan 1853) [See Sherborn, 1934]. 


Grose-Smith, Henley (1833-1911), and William Forsell Kirby (1844-1912) 


1887-1902. Rhopalocera exotica, being illustrations of new, rare, and unfigured species 
of butterflies. London: Gurney & Jackson. 3 vols. [Exact dates for different sections 
need to be ascertained from the appropriate species name, since the volumes are 
completely mixed as to dates for the various species and pages are not consecutive 
but involve only each species description in most cases.] 


Guenée, Achille (1809-1880) 


1845. Essai sur une nouvelle classification des microlépidopteres et catalogue des 
especes Européenes connues jusqu’a ce jour. Ann. Soc. Ent. France (2) 3:105-192, 
297-344. 


[1846]. Europaeorum microlepidopterorum index methodicus. Paris: Roret. 106 pp. 
[Reprint of 1845 paper]. 
Guérin-Méneville, Felix Edouard (1799-1874) 


1830-[1838]. Crustacés, arachnides et insectes. In Voyage autour du monde ... sur 
eeawoquille pendant 4). 1622-25 ..> par M. lL. 1. Dupemy, & c. Vol. 2-(2) (1). 
Paris. 319 pp., 22 pls. (Lepidoptera: pp. 271-286 [10 Dec 1838]) [See Cowan, 197 la]. 


1830 (26 May) Piselo—14 
1831 (29 May) aE 
1830 ( 1 Sep) 15-16 
(25 Nov) 17 
1831 (15 Jun) 18 
( 7 Mar) 19 


1829-1844. Rhopalocera. In Iconographie du regne animal de G. Cuvier, ou repreé- 
sentation d’apres nature de l'une des éspeces les plus remarquables et souvent non 
figurées de chaque genre d’animaux. Avec un texte descriptif mis au courant de la 
science. Ouvrage pouvant servir d’atlas a tous les traités de zoologie. Paris. (Insects: 
576 pp., 110 pls.) [See Cowan, 1971d]. 


1829 Pls. 3-12 

1830 13-14, 21-24 

1831 25-30, 42-46, 52-55, 60, 62, 76, 24-25 bis, 28 bis 
1832 77-92, 99, 84 bis 

1833 31, 36-39, 47-49, 59, 101, 39 bis 

1834 32-35, 50, 61, 64-66, 49 bis 

1835 15-20, 40, 51, 56-58, 67-75, 93-98 

1836 100, 102-104 

1837 1-2, 41, 63 


1844 Text pages 1-576 


96 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Haworth, Adrian Hardy (1767-1833) 


1803-1828. Lepidoptera Britannica, sistens digestimen novam lepidopterorum quae 


in Magna Britannica reperiunter ... adjunguntur dissertationes variae ad historiam 
naturalam spectantes. London: J. Murray. 609 pp. [4 parts]. 

1803 [Part 1] Pp. 1-136 

[1809] [Part 2] 137-376 

[1811] [Part 3] 377-512 

1828 [Part 4] 513-609 


Heinemann, Hermann von (1812-1871) 


1863-[1876]. Die Schmetterlinge Deutschlands und der Schweiz. Zweite Abthei- 
lung. Kleinschmetterlinge. Braunschweig: Schwetschke. 2 vols. [With F. Wocke for 
Vol. 2, part 2]. 


1863 Vol. 1 (Part 1) Pp. 1-248; plus 1-39 
1865 Vol. 1 (Part 2) 1-214; plus 1-27 
1870 Vol. 2 (Part 1) 1-388 

[1876] Vol. 2 (Part 2) 389-825; plus 1-102 


Herrich-Schaffer, Gottlieb August Wilhelm (1799-1874) 


1843-1856. Systematische Bearbeitung der Schmetterlinge von Europa, zugleich als 
Text, Revision und Supplement zu Jakob Hubner’s Sammlung europaischer Schmet- 
terlinge. Regensburg: G. J. Manz. 6 vols. [See Hemming, 1937]. 


Volume 1: Die Tagfalter 


1843 Papilionides Pp. 1-40 Pls. 1-5, 29-34 
1844 Al—122 6-28, 35-52 
1845 123-162 53—57 
1846 163-164 58-70 
1847 = 71-77 
1848 —_ 78-79 
1849 — 90-91 
1850 — 92-102 
1851 — 103-118 
1852 = 119-127, 130-132, 134 
1853 = 128-129, 133 
1854 [Index] 120 
1855 [Index] pA! 
1845 Hesperides — Pls. 1-3 
1846 a 4-5 
1848 a 6 
1854 -= i 

Volume 2: Die Schwarmer, Spinner und Eulen 
1846 Text Pp. 1-104 
1847 105-166 
1848 167-190 
1849 191-238 
1850 239-334 
1851 335-450 
1855 [Index] 1-64 
1844 Plates: Hepialides Pls. 1 
1846  Hepialides and Cossides 1 
1851 2 
1843 Zygaenides 1-2 
1844 3-7 
1846 8-11 
1847 1Q=13 


VOLUME 36, NUMBER 2 Q7 


1851 14-16 
1846 — Sesiides l=s 
1848 9 
1851 10 
1843 Sphingides IL 
1844 2 
1847 3-4 
1843  Bombycides =) 
1844 3-9 
1846 10-16 
1847 Ni BY 
1848 Dei 
1850 28 
1851 29-31 
1853 32 
1843 Noctuides 13} 
1844 4-8 
1845 9-23 
1846 94-52 
1847 53-69 
1849 70-82, 85-87 
1850 83-84, 88-89, 101-102 
1851 100, 103-113 
1852 114-124 
1853  Nycteolidae 1 
Volume 3: Die Spanner 
1843 Geometrides Pls. 1 
1844 Bp, 123 2-11 
1845 _— 12-20 
1846 9-16 LAX) 
1847 a2 41-56 
1848 73-[184] S771 
1850 = Pe 
1851 —_ 75-87 
1853 _ 88 
1854 = 89-90 
1855 [Index] 134: 
Volume 4: Die Zunsler und Wickler 
Pyralidides Tortricides 
1847 Text Pp. — Pls. 14 Pls. 1-20 
1848 1-80 5-10 QA 
1849 81-128 LY 48-52 
1850 as aall, 53-54 
1851 129-288 18-20 = 
1852 = Piles 56-57, 59 
1853 a = 55 
1854 = = 58 
1855 [Index] 1-48 
Volume 5: Die Schaben und Federmotten 
1847 Text Pp. — Tineides Pls. 1-10 
1848 a es 
1849 = 19-23 
1850 — 24-36, 38, 40-41 
1851 = 7, BO), 28), Bi’ SOAS 
1852 = 54, 56, 96-103 
1853 72 55, 58, 63-74, 80-88, 94-95, 


105-108, 116 


98 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


1854 73-224 75-79, 89-93, 104, 109-115, 
117-124 
1855 225-394 ; index 1-52 
1850 Fterophides 1-2 
1851 Micropteryges 1 
1853 Pterophides 4-6 
1854 Pterophides 3,7 
Volume 6: Sechster und letzter Band 
1843 Text Pp. — Pls. Macros: 1 Micros: — 
1844 [Macros] 1-6 2-11 — 
1845 — 2 aus 
1846 11-14 13-14, 16 == 
1847 — — 1-3 
1848 = 15, 17-19 — 
1849 15-18 20=22 4-9 
1849 [Micros] 14 
1853 — — 10-14 
“Nachtrag zum ersten Bande”’ 
1851 Pp. l=24 
1852 25-80 
1855 81-152 
1856 153-178 
“Systema Lepidopterorum Europae”’ 
1852 Pp. 1-36 
1853 37-40 
1854 41-60 
1855 ol=2 
1856 [Index] 1-48 


1850-[1869]. Sammlung neuer oder wenig bekannter aussereuropaischer Schmet- 
terlinge. Regensburg. 2 vols. [See Dalla Torre, 1927; Fletcher, 1979]. 


Volume 1. Series 1: Heterocera (Nachtfalter) 


[1853] Pls. 1-16 
[1854] 17-48 
[1855] 49-68 
1855 lias 
[1856] 80-91 
[1858] 69-70, 79, 95-96 
Volume 2. Series 2: Rhopalocera (Tagfalter) 
1850 Pls. 1-10 
[1853] Wane! 
[1855] 15-18 
[1856] Pp. t=52 19222) 
[1858] 53-84 23-24 
Volume 2. Series 1: Heterocera (cont.) 
[1869] 97-100 
Volume 2. Series 2: Rhopalocera (cont.) 
[1869] Pp... 14 25-28 


[1856-1861]. Neue Schmetterlinge aus Europa und den angrenzenden Landern. 
Regensburg. 32 pp., 26 pls. [Pages 21-23 are missing through a typographical error]. 


[1856] Pp. 1-8 Pls. 1-9 
1860 9-20 10-18 
[1861] 24-32 19-26 


Hewitson, William Chapman (1806-1878) 


1852-1877. Exotic butterflies, being illustrations of new species selected chiefly from 
the collection of William Wilson Saunders and William C. Hewitson. London: V. 


VOLUME 36, NUMBER 2 99 


Voorst. 3 vols. [See Griffin, 1932b, for dates of each plate; corrections of some 
plates in Hemming, 1945]. 


1863-1878. Illustrations of diurnal Lepidoptera. London: V. Voorst. 229 pp., 90 pls. 
[Plates 86-87 are missing through a typographical error]. [See Hemming, 1935]. 


1863 Pp. 1-36 Pls. 1-16 

1865 37-76 17-30 

1867 77-114 31-46 

1869 115-136, 14a-h, Suppl. 1-16 47-54, 3a—c, Suppl. I-V 
1873 137-150 55-59, Suppl. VI 

1874 151-184 60-73 

1877 185-208 74-83 

1878 209-229, Suppl. 35-47 84-85, 88-92, Suppl. 


la—b.iilia—b. Nas 
Va-b, VII, VIII 


Hubner, Jacob (1761-1826) 


1796-[1838]. Sammlung europaischer Schmetterlinge. Augsburg. 7 vols. [9 parts]. 
[See Hemming, 1937]. 


[Part] I: Papilios 


[1799-1800] Pisaeel—soo 
[1800-03] 89-96 

_ [1803-04] 97-114 
[1805] UE IUUS) 
[1805-06] 120-124 
[1806-08] 125-128 
[1808-13] 129-144 
[1814-16] 145-148 
[1816-17] 149-150 
[1818-19] Lit 
[1819-22] SeQ= 54 
[1822-23] 155 
[1823] 156-161 
[1823-24] 162-175 
[1824-25] 176 
[1825-26] ISIS I 
[1827-28] 182-187 
[1828-32] 188-195 
[1832-33] 196 
[1834-36] 197-201 
[1836-38] 202-203 
[1837-38] 204-207 

[Part] II: Sphinges 
1796 is, LENS 
[1803-06] 1 (rev.) 
[1796-99] Me, 
[1803-06] 18-22 
[1807-08] ye} 
[1808-13] 24-29 
[1814-17] 30 
[1818-19] Sill 
[1819-22] O22 54 
i425] 35 
[1827-28] 36 
[1834-36] Sivi 
[1836-38] 38 


100 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


[Part] III: Bombyces 


[1800-03] Pls. 1-43 
[1803-08] 44-57 
[1809-10] 58-59 
[1810-13] 60-63 
[1818-19] 64-65 
[1819-22] 66 
[1823] 67 
[1823-24] 68-69 
[1827] 70 
[1827-28] 71-74 
[1828-29] Te T6 
[1830-31] 77-80 
[1834-36] 81 
[1836] 82 
[1836-38] 83 
[Part] IV: Noctuae 
[1800-03] Pls..eHl=74 
[1803-08] 75-86 
[1808-09] 87-96 
[1809-13] 97-134 
[1814-17] 135-139 
[1818-19] 140-141 
[1819-22] 142-150 
[1823] 151 
[1823-24] 152-156 
[1825-26] 157 
[1827-28] 158-160 
[1828-30] 161-162 
[1828-32] 163-169 
[1832-33] SO = el 
[1834] 172-176 
[1834-35] 177-180 
[1836] 181 
[Part] V: Geometrae 
[1796-99] Pls. 1-59 
[1799-1800] 60 
[1800-09] 61-66 
[1809-13] 67-89 
Msi] 90-95 
[1818-19] 96-98 
[1819-22] 99-100 
[1824-25] 101-102 
[1825-26] 103-105 
[1828-31] 106-108 
[1832-33] 109-111 
[1838] Leos 
[Part] VI: Pyralides 
1796 Piso ael=20 
[1796-99] EAL 
[1800-09] 22-23 
[1811-13] 24-27 
[1818-19] | 28-29 
[1823] 30 
[1828-32] 31 


[1832-33] 32 


VOLUME 36, NUMBER 2 101 


[Part] VII: Tortrices 


[1796-99] Pilss v9) 
[1799-1800] 30 
Sia = 13) Sul 8h7/ 
[1814-17] 38-41 
[1818-19] 42-43 
[1819-22] 44 
[1822-23] 45 
[1823] 46 
[1823-24] 47 
[1830] 48-52 
[1832-33] 53 
[Part] VIII: Tineae 
1796 Pils, 84! 
[1796-99] 50-51 
[1800-05] 38-42 
[1805-10] 43-44 
[1810-13] 45-63 
[1814-17] 64-66 
[1818-19] 67 
[1823-24] 68-69 
[1832] 70 
[1834-36] Tal 
[Part] IX: Alucitae 

[1800-05] Pils. lee 
[1805-13] 3-6 
[1818-19] a 


[1806]. Tentamen determinationis digestionis atque denominationis singularum stir- 
pium lepidopterorum, peritis as inspiciendum et dijudicandum communicatum, a 
Jacobo Hubner. 2 pp. [Rejected: Opinion 97 (ICZN, 1926)]. 


1806-[1838]. Sammlung exotischer Schmetterlinge. Augsburg. 3 vols. [Plates are un- 
numbered: see Hemming, 1937, for exact plate dates]. 


1806-[1819] Volume 1 
[1819-27] Volume 2 
[1827-38] Volume 3 


1816-{1826]. Verzeichniss bekannter Schmettlinge [sic]. Augsburg. 431 pp.; plus 
72 pp. 


1816 Pp 16 
[1819] 17=176 
[1820] 177-208 
[1821] 209-256 
[1823] 257-304 
[1825] 305-431 
“Anzeiger” 
[1826] 72, 


Illiger, Johann Car! Wilhelm (1775-1813) 


1807. Die neueste Gattungs-Eintheilung der Schmetterlinge aus den Linneischen 
Gattungen Papilio und Sphinx. Magazin fur Insektenkunde 6:227-295. [Opinion 232 
(ICZN, 1954): new genera authored by Latreille and Fabricius, not Illiger]. 


Kluk, Krzysztof (1739-1796) 


1780. Zwier at domowych i dzikich osobliwie krajowych .... Warsaw. [See Paclt, 
1955, for details]. 


102 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Latreille, Pierre André (1762-1833) 


1809-1817. Les insectes. In F. H. H. von Humboldt and A. J. A. Bonpland, Voyage 
aux régions €quinoxiales du nouveau continent, fait en 1799-1804, & c. Pt. II. [Zo- 
ology]. Recueil d’observations de zoologie et l’anatomie comparée; faites dans 
VYocéan Atlantique, dans l’intérieur du nouveau continent, & c. Paris. [See Mac- 


Gillavry, 1931]. 


1809 Vol. 1 [Pt. 9 of vol. 1 of Zool.] Pp. 197-283 Pls. 15-18 
344-397 2225 

1813 Vol. 2 [Pt. 2 of vol. 2 of Zool.] 65-96 — 

1817 97-138 31-43 


Lucas, Pierre Hippolyte (1815-1899) 


1857. Lepidoptera. In R. de la Sagra, Histoire physique, politique et naturelle de 
V’Ile de Cuba, & c. Paris. Vol. 7:202-313. [See Aguayo, 1946. Spanish edition also 
dated 1857]. 


Ménétriés, Eduard (1802-1861) 


1855-1863. Enumeratio corporum animalium musei imperialis academiae scientia- 
rum Petropolitanae. Classis insectorum, ordo lepidopterorum. Petropolis [=Lenin- 
grad]. 3 vols. 


1855 Volume 1 Pp. 1-66 Pls. 1-6 
[plus pp. 67-97] 

1857 Volume 2 Pp. 67-112 Pls. 7-14 
[plus pp. 99-144] 

1863 Volume 3 Pp. 1-20 Pls. 1-4 


Mikan, Johann Christian (1769-1844) 
1820-1825. Delectus florae et faunae brasiliensis. Vienna. [See Stearn, 1956]. 


1820 Part 1: Pp. 1-6 Pls. 1-6 

1822 Part 2: 1-6 7-12 
1823 Part 3: 1-6 13-18 
1825 Part 4: 1-6 19-24 


Moore, Frederic (1830-1907) 
1880-1887. The Lepidoptera of Ceylon. London: Reeve. 3 vols. [See Griffin, 


1939}. 
1880 Volume 1: Pp. 1-40 Pls. 1-18 
[1881] 41-190 19-71 
1882 Volume 2: 72 (207 
[1883] Toa-No2, 108-143 
1884 Volume 3: 1-88 144-157 
[1885] 89-304 158-181 
[1886] 305-392 182-195 
1887 393-578 196-215 


Perty, Joseph Anton Maximilian (1804-1884) 
1833. Lepidoptera. In Delectus animalium articulatorum quae in itinere per brasi- 
liam.... Munich. Pp. 151-164; pls. 29-32. (1830-34). 
Rambur, Jules Pierre (1801-1870) 
1837-1842. Faune entomologique de |’ Andalousie. Paris: A. Bertrand. 2 vols.; 19 pls. 
1837 Volume 1: Pp. 41-80 


1838 (Mar) 81-144 
Volume 2: 16 


VOLUME 36, NUMBER 2 103 


1838 (Dec) 17-96 

1839 (Jan) 97-176 

1840 (Mar) 177-304 

1842 305-336 [incomplete] 
1842 Reprint (336 pp.) 

1942 Facsimile reprint. Madrid. 


1858-{1866]. Catalogue systematique des lepidopteres de lAndalousie. Paris: 
Bailliere. 442 pp. 


1858 Pp. 1-92 
[1866] 93-442, 
Seitz, Adalbert (1860-1938) [Editor] 


1906-1954. Die Gross-Schmetterlinge der Erde. Stuttgart: A. Kernen. 16 vols. [See 
Griffin, 1936; Nye, 1975]. (The Macrolepidoptera of the World.) 


Vol. 5. Die amerikanischen Tagfalter. (The American Rhopalocera). 203 pls. 
English Edition German Edition 


1907 Ep: 1-40 Pp. 1-48 

1908 41-48 49-72 

1909 49-160 73-160 

1910 161-200 161-200 

1911 201-240 201-240 

1912 241-344 241-344 

1913 345-448 345-456 

1914 449-480 457-480 

1915 481-536 48 1-536 

1916 537-656 537-664 

LEST 657-720, 857-864 665-720 

1920 721-744 721-744 

VIS NS) 745-768 745-768 

1920 769-8 16 769-8 16 

1921 817-856 817-856 

1922 865-904 857-912, 937-944 
1923 905-928 913-936, 945-1000 
1924 929-1132 1001-1141 


Vol. 6. Die amerikanischen Spinner und Schwamer. (The American Bombyces and 
Sphinges.). 186 pls. (185 pls. English ed.). 


English Edition German Edition 


1913 Pp. 1-32 Pp. 1-32 
1915 33-192 33-192 
1917 193-232 193-240 
1918 233-280 249-280 
1919 28 1-320 241-248, 281-320 
1920 321-336 321-336 
1921 337-376 337-376 
1922 377-416 377-416 
1924 — 417-424 
1925 417-528 425-528 
1927 529-616 529-616 
1928 617-672 617-672 
1929 673-768 673-768 
1930 769-840 769-840 
1931 841-904 841-904 
1932 905-1016 905-1016 
1933 1017-1048 1017-1048 


1934 1049-1088 1049-1088 


104 


1935 
1936 
1937 
1938 
1939 
1940 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


1089-1112 
1137-1256 
1113-1136, 1257-1296 
1297-1304 


1089-1112 
1137-1256 
1113-1136, 1257-1296 
1297-1304 
1305-1328 
1329-1452 


Vol. 7. Die Gross-Schmetterlinge des amerikanischen Faunengebietes. Eulenartige 
Nachtfalter. (The Macrolepidoptera of the American Region. Noctuiformes). 96 pls. 


English Edition 


German Edition 


1919 Ep 1=12 Pp. 1-20 
1924 13-156 21-180 
1925 157-228 181-228 
1926 229-308 229-308 
1927 309-348 309-348 
1928 349-364 349-364 
1930 365-380 365-380 
1936 381-396 381-396 
1939 397-412 [incomplete] 397-428 
64 pls. 
1940 429-468 
1944 469-508 [incomplete] 
96 pls. 


Vol. 8. Die amerikanischen Spanner. 17 pls. 
German Edition 


1932 i Pp. 1-56 

1933 Sy 57-712 

1935 Hi, 73-80 

1936 ai 89-104 

1938 Gu 81-88, 105-144 
[incomplete] 


Sepp, Jan (1778-1853) 


1838-1852. Natuurlijke Historie van Surinaamsche Vlinders, qawnlherlemen geteek- 
end. Amsterdam. 328 pp. [3 pts.], 152 pls. 


1828-{1848] [Part 1]: Pp. 1-108 Pls. 1-50 
[1832-40] [Part 2]: 109-224 51-100 
[1852]-1855 [Part 3]: 225-328 101-152 


Staudinger, Otto (1830-1900) and Ernst Schatz (1844-1887) 


1884-1892. Exotische Tagfalter in systematischer Reihenfolge mit Berucksichti- 
gung neuer Arten. Furth: G. Lowensohn. 2 vols., 150 pls. [New taxa by Stau- 
dinger]. 


1884 Volume 1: Pp. 1-38 Pls. 1-30 
1885 39-102 31-60 
1886 103-174 61-80 
1887 175-234 81-95 
1888 235-333 96-100 
1885 Volume 2: Pp. 1-32 1-10 
1886 33-92 11-16 
1887 93-136 17-26 
1888 137-180 27-34 
1889 181-224 35-42 
1892 225-284 43-50 
Stephens, James Francis (1792-1852) 
1827-1835. Illustrations of British entomology; or, a synopsis of indigenous in- 


sects; containing their generic and specific distinctions; with an account of their 


VOLUME 36, NUMBER 2 105 


metamorphoses, times of appearance, localities, food, and economy, as far as prac- 
ticable. Haustellata. London: Baldwin and Cradock. 4 vols., 41 pls. 


1827 Volume 1: Pp. 1-56 Pls. © 1-9 
1828 57=152 10-12 
1828 Volume 2: 1-80 13-18 
1829 81-203 19-24 
1829 Volume 3: 1-96 DESY 
1830 97-136 28=sil 
1831 ISTASOy 32 
1831 Volume 4: _— 33-41 
1834 1-352 = 
1835 353-436 — 


1829a. The nomenclature of British insects; being a compendious list of such species 
as are contained in the systematic catalogue of British insects, and forming a guide 
to their classification, & c. & c. London: Baldwin and Cradock. 68 pp. [June 1, 1829]. 


1829b. A systematic catalogue of British insects. London: Baldwin and Cradock. 2 
parts. 804 pp. [July 15, 1829]. 


Stoll, Casper (?—1795) 
1787-1790. Aanhangsel van het Werk, de Uitlandsche Kapellen, voorkomende in de 
drie Waereld-Deelen Asia, Africa en America, door den Heere Pieter Cramer, ver- 
vattende naauwkeurige afbeeldingen van Surinaamsche Rupsen en Poppen; als mede 
van veele zeldzaame en nieuwe ontdekte Uitlandsche Dag- en Nagt-Kapellen. Am- 
sterdam: Gravius. 184 pp., 42 pls. [“Volume 5” of Cramer]. 


1787 Pp. 1-42 Piss 27 
1790 43-184 8-42 


Strand, Embrik (1876-1947) 


1914-1918. Lepidoptera Niepeltiana, Abbildungen und Beschreibungen neuer und 
wenig bekannter Lepidoptera aus der Sammlung W. Niepelt. Leipzig: Urban. 
94 pp., 18 pls. 


1914 [Part 1]: Pp. 1-64 Pls. 1-12 
1916 [Part 2]: 1-26 13-17 
1918 [Part 3]: 1-4 18 


Strecker, Ferdinand Heinrich Hermann (1836-1901) 


1872-1900. Lepidoptera, Rhopalocera and Heteroceres, indigenous and exotic; 
with descriptions and colored illustrations. Reading, Pennsylvania. 15 parts; 3 suppl.; 
15 pls. [See Brown, 1964a; Griffin, 1931b; Oiticica, 1946]. 


1872 Part 1: Pp. 1-8 Pls; tal 
1873 ete, Moe 9-60 2-7 
1874 Pt. 8-11: 61-100 8-11 
1875 Pt, 12: 101-108 12 
1876 Pye 1833 109-123 13 
1877 Pt. 14-15: 124-143 14-15 
1898 Suppl. 1: 1-12 — 
1899 Suppl. 2: 1-11 — 
1900 Suppl. 3: 13-37 — 


Swainson, William (1789-1855) 
1820-1823. Zoological illustrations, or original figures and descriptions of new, 
rare or interesting animals... . (Series 1). 


1820 Part 1: Pils, Wes 
1821 19-66 


106 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


1821 Part 2: 67-83 
1822 84-119 
1822 Part 3: 120-134 
1823 135-182 
1829-1833. Ibid. (Series 2). 
1829 Part 1: Pls. 1-30 
1830 31-45 
1831 Part 2: 46-85 
1832 86-91 
1832 Part 3: 92-96 
1833 97-136 


Walker, Francis (1809-1874) 


1854-1866. List of the specimens of lepidopterous insects in the collection of the 
British Museum. London. 35 parts. [See Nye, 1975]. 


1854 Part 1: (Heterocera) Pp. 12278 
B Pare: 279-58 1 
1855 Part 3: 582-775 
f Part 4: 776-976 
fi Part 5: 977-1258 
d Part 6: 1259-1508 
1856 Part 7: 1509-1808 
iD Part 8: (Sphingidae) 1-271 
v Part 9: (Noctuidae) 1252) 
[1857] Part 10: 253-492 
1857 Part 11: 493-764 
[1858] Part 12: 765-982 
4 Part 13: 983-1236 
1858 Part 14: 1237-1520 
‘ Part 15: 1521-1888 
[1859] Part 16: (Deltoides) 1-254 
1859 Part 17: (Pyralides) 255-508 
i Part 18: 509-798 
fh Part 19: 799-1036 
1860 Part 20: (Geometrites) 1-276 
: Part 21: 277-498 
1861 Part 22: 499-756 
Part 23: 757-1020 
1862 Part 24: 1021-1280 
if Part 25: 1281-1478 
[1863] Part 26: 1479-1796 
1863 Part 27: (Crambites & 1-286 
4 Part 28: Tortricites) 287-562 
1864 Part 29: (Tineites) 563-836 
i Part 30: 837-1096 
[1865] Part 31: (Supplement) 1322 
1865 Part o2: 323-706 
U Part 33: 707-1120 
[1866] Part 34: 1121-1534 
1866 Part 35: 1535-2040 


Walsingham, Thomas de Grey (1843-1919) 


1892. On the micro-lepidoptera of the West Indies. Proc. Zool. Soc. Lond. 1891: 
492-548; pl. 41. [not 1891]. 


VOLUME 36, NUMBER 2 107 


1908. Microlepidoptera of Tenerife. Proc. Zool. Soc. Lond. 1907:911-1034; pls. 
51-53. [not 1907]. 

1909-1915. Tineina, Pterophorina, Orneodina and Pyralidina and Hepialina (part). 
In F. D. Godman and O. Salvin, Biologia Centrali-Americana. [Vol. 42]. Insecta: 
Lepidoptera-Heterocera. Vol. 4. London. 482 pp.; 9 pls. 


1909 Pp. 1-24 

1910 25-48 

1911 49-112 

1912 113-168 

1913 169-224 

1914 225-392, 

1915 393-482 9 pls. [undated] 


Westwood, John Obadiah (1805-1893) 


[1838]-1840. Synopsis of the genera of British insects. London. 158 pp. [published 
in Vol. 2 of work cited below]. 


[1838] Pp. 1-48 
[1839] 49-80 
1840 81-158 


1838-1840. An introduction to the modern classification of insects; founded on 
natural habits and corresponding organization of the different families. London: 
Longman. 2 vols. [Dates after Griffin, 1932.] 


1838 Vol. 1: Pp. 1-462 
1839 Vol. 2: Pp. 1-352 
1840 353-587 


1843-1845. In H. N. Humphreys and J. O. Westwood, British moths and their 
transformations. London: Smith. 2 vols. 


1843 Vol. 1 
1845 Vol. 2 


Wood, William (1774-1857) 


1833-1839. Index entomologicus; or, a complete illustrated catalogue, consisting 
of 1944 figures, of the lepidopterous insects of Great Britain. London. 266 pp.; 


54 pls. 


1833 Poe 232 Ps. 1) 
1834 33-88 10-19 
1835 89-128 20-29 
1836 129-168 30-38 
1837 169-214 39-47 
1838 215-266 48-54 
1839 Title page & preface 


Zeller, Phillip Christoph (1808-1883) 


1852. Lepidoptera Microptera, quae J. A. Wahlberg in Caffrorum terra collegit. 
Glogau. 120 pp. [Reissued in [1854] under the same title in Kongl. Svenska Vetensk. 
Akad. Handl. 1852:1-120.] 


Zetterstedt, Johan Wilhelm (1785-1874) 


[1839]-1840. Insecta Lapponica. Lipsiae: Voss. 1140 pp. 
[1839] Pp. 1-1014 
1840 1015-1140 


108 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


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Smithsonian Misc. Coll. 73: 19-30. 


110 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


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1957b. Opinion 474. Determination of the dates to be assigned for the pur- 

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1958. Opinion 516. Determination under the plenary powers of the relative 
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VOLUME 36, NUMBER 2 IEA 


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Journal of the Lepidopterists’ Society 
36(2), 1982, 112-118 


THE CHROMOSOMES OF A WILD SILKMOTH, 
ARCHAEOATTACUS EDWARDSII, WITH A RECORD 
HIGH CHROMOSOME NUMBER 
FOR SATURNITDAE 


R. C. NARANG! AND M. L. GUPTA? 


ABSTRACT. The diploid chromosome number of Archaeoattacus edwardsii (White), 
new to chromosome cytology, is 128, the highest so far reported in the family. The 
karyotype is characterised by the presence of two exceptionally larger (probably sex) 
chromosomes. The sex mechanism is XX ¢ : XY 2. The females show a prominent sex 
chromatin body in germ as well as somatic cells. The evolutionary status of the species 
has been discussed. 


Besides the usually high number and smaller size of the chromo- 
somes, the cytogenetical studies in moths also involve difficulties re- 
garding the procurement of stages from caterpillar to adult, as the 
mitotic and the male meiotic divisions are passed before the adult 
stage. So far, the chromosome numbers of only 30 saturniid species 
(including the present report), belonging to 16 genera (Table 1) out 
of about 1000 species recorded, are known. Interestingly, the chro- 
mosome number in Archaeoattacus edwardsii (White) is the highest. 
The karyotype and the sex chromatin in this species are described 
here for the first time. 


MATERIAL AND METHODS 


For the present study the cocoons were collected from Khasi Hills 
(Meghalaya) during Sept.—Oct. 1979. The slides were prepared from 
the gonads and brain tissues of both sexes. Some cocoons were also 
raised to the adult stage for the systematic determination of the species. 
Giemsa-stained cytological preparations were made by heat-dry smear 
technique (Narang & Gupta, 1979a), using, however, a 5% concentra- 
tion of the stain. Some of the preparations were made after injection 
of 0.1 ml of 0.05% colchicine for 2.30 to 3 h. 


OBSERVATIONS 


The diploid number of chromosomes has invariably been found to 
be 128 at mitotic metaphase in germinal (Figs. 1 & 2) as well as 
neuroblast cells (Fig. 3) in as many as 15 cells of 6 males and 20 cells 
of 3 females. The same chromosome number has been confirmed by 
the count of 64 bivalents in 5 male diakinetic cells (Fig. 4). The karyo- 


' Department of Zoology, M.M.(P.G.) College, Modinagar-201 204 INDIA. 
2 Department of Zoology, University of Jodhpur, Jodhpur-342 001 INDIA. 


VOLUME 36, NUMBER 2 


o,* co 
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Fics. 1-5. Chromosomes of Archaeoattacus edwardsii: 1, spermatogonial meta- 
phase; 2, oogonial metaphase from colchicinized individual showing localized centro- 
meres; 3, mitotic metaphase from neuroblast cell (¢); 4, diakinesis (¢); 5, oogonial 
interphase showing sex chromatin (SC). Arrows indicate larger (sex ?) chromosomes. 


Bar represents 10 um. 


114 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


type is further characterized by the presence of two exceptionally 
larger chromosomes. The average length of these chromosomes is 
nearly the same (i.e., 2.5 wm in male and 2.5 & 2.7 um in female). 
The smaller chromosomes range in length from 0.5 wm to 1.25 wm. 

The mitotic metaphase chromosomes of edwardstii have been found 
to be of monocentric type as studied in the colchicinized preparations. 
Fig. 2 reveals an interstitial centromere in some of the chromosomes; 
whereas, the position of the centromere in others is not discernible, 
due to their smaller sizes. 

A prominent, large-sized and positively heteropycnotic sex chro- 
matin body has been clearly observed in interphase and prophase 
cells of brain and germ tissues of the females. The average size of sex 
chromatin at the interphase stage (Fig. 5) is 1.15 um. 


DISCUSSION 


The presence of the same and even number of the chromosomes 
in both the sexes and the formation of a sex chromatin body in the 
females indicates the sex chromosome mechanism in Archaeoattacus 
edwardsii to be XX 6: XY 2, as concluded for eight other saturniid 
species (Gupta & Narang, 1980a; Gupta & Narang, 1981); the sex 
chromatin, formed both in germ and somatic cells, represents the 
Y-chromosome. 

Bauer (1943) and White (1957) suggested that the big chromosome 
frequently found in Lepidoptera is a sex chromosome, at least in those 
species which have a very high chromosome number. In the family 
Saturniidae to which the present species belongs, the modal chro- 
mosome number is n = 31, which has been reported in 18 species out 
of the 30 investigated so far (Table 1). The two exceptionally larger 
chromosomes of this species, with n = 64, seem to, likewise, repre- 
sent the sex chromosomes. The difference in the size of these chro- 
mosomes compared to the autosomes is probably because the sex 
chromosome mechanism in this group is nicely balanced and fission 
or other such rearrangements in sex chromosomes may disturb the 
mechanism. This is also clear from the fact that there are very few 
cases in Lepidoptera in which multiple sex chromosomes are known. 

In the tribe Attacini of the subfamily Saturniinae, seven species, 
four from genus Hyalophora and one each from Philosamia, Callo- 
samia and Archaeoattacus, have so far been cytogenetically worked 
out. The haploid chromosome number in all the species of Hyalo- 
phora is 31, while it is 13-14 in Philosamia cynthia (Drury) (in its 
different races), 19 in Callosamia promethea (Drury) and 64 in A. 
edwardsii. Further work in other genera and species of Attacini is 
urgently needed for exploring the evolutionary trends in this tribe. 


VOLUME 36, NUMBER 2 als) 


TABLE 1. Known haploid chromosome numbers for species of Saturniidae. 


Haploid 
chromosome References 
S. No. Name of the species number (First report) 
SUBFAMILY: SATURNIINAE 
TRIBE: SATURNIINI 
i Actias selene 31 Deodikar et al. (1969) 
2. A. luna 31 Unpublished 
3. Antheraea assamensis 15 Deodikar et al. (1962) 
A. A. compta 15 Gupta & Narang (1981) 
5: A. frithi 31 Jolly et al. (1977) 
6. A. mylitta 31 Sinha & Jolly (1967) 
ae A. pernyi 49 Kawaguchi (1933, 34) 
8. A. (=Telea) polyphemus 30 Cook (1910) 
9. A. roylei Sl Jolly et al. (1970) 
10. A. sivalica Sul Jolly et al. (1978) 
alk A. yamamai 31 Kawaguchi (1933) 
1 Cricula trifenestrata Sil Narang & Gupta (1979a) 
{3: Dictyoploca cachara 30 Narang & Gupta (1979b) 
14. D. japonica Sill Oba (1942) (cited by 
Makino, 1951) 
15! D. simla oll Unpublished 
16. Graellsia isabelae Sill Templado et al. (1975) 
ike Loepa katinka 28 Narang & Gupta (1979c) 
18. Saturnia pyri 30 Pariser (1927) 
1.2: Sonthonnaxia maenas 3] Narang & Gupta (1979d) 
20. Eriogyna pyretorum 30 Gupta & Narang (1980b) 
2, Eudia (=Saturnia) pavonia 29 Kernewitz (1915) 
TRIBE: ATTACINI 
O2.. Archaeoattacus edwardsii 64 (Present work) 
23) Philosamia cynthia 13-14 Dederer (1907, 15); 
Deodikar & Thakar 
(1958) 
24. Hyalophora (=Platysamia) cecropia 31 Bytinski-Salz (1938) 
25. H. euryalis ol Bytinski-Salz (1938) 
26. H. gloveri ol Bytinski-Salz (1938) 
OT. H. columbia 31 Bytinski-Salz (1938) 
28. Callosamia promethea 19 Cook (1910) 
SUBFAMILY: HEMILEUCINAE 
TRIBE: HEMILEUCINI 
29. Automeris io ol Cook (1910) 
SUBFAMILY: CITHERONIINAE 
30. Anisota (=Dryocampa) rubicunda Sul Ennis (1976) 


In the sister tribe Saturniini the probable modal number as indicated 
by the available data (Table 1) is n = 31, which is also the modal 
number for Lepidoptera as a whole (Suomalainen, 1969; White, 1973). 
From this it can be assumed that the same number, n = 31 (at present 
known only from genus Hyalophora), might be the ancestral number 


116 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


of the tribe Attacini, and the lower chromosome number of Philosa- 
mia cynthia (n = 13-14) and Callosamia promethea (n = 19), and the 
higher number for A. edwardsii (n = 64) might have evolved from 
this number by the mechanism of chromosomal fusions and fissions 
respectively (see discussion below). The presence of a single pair of 
exceptionally larger (probably sex) chromosomes and simple XX 6 : XY 
2 sex mechanism in A. edwardsii speaks against an incidence of poly- 
ploidy. 

Bigger (1975, 76) clearly showed localized centromeres in the mi- 
totic chromosomes of five butterflies and pointed out that there are 
two types of centromeric organizations. According to him at early mi- 
totic metaphase the chromosomes exhibit a monocentric organization, 
either exclusively or, more probably, with one portion of the diffuse 
centromere exerting a dominant influence over the remainder. He 
further stated that with the advancement of metaphase the influence 
of this centromere is either lost or superceded by the combined in- 
fluence of the rest of the centromere. Several other authors (e.g., Dan- 
ilova, 1973; Gassner & Klemetson, 1974; Rishi & Rishi, 1979; Gupta 
& Narang, 1981) have also reported clear primary constrictions (lo- 
calized centromere) at mitotic and/or meiotic II metaphases in several 
Lepidoptera. But there has appeared absolutely no report, either by 
light microscopy or by electron microscopy, about the localized nature 
of the centromere at metaphase I in this group. This dual centromeric 
organization and the presence of morphologically distinguishable sex 
chromosomes in Lepidoptera probably explains why the chromosome 
numbers in this group are less variable than the diffuse kinetochore 
(e.g., in Luzula) would at least theoretically allow. 

It is a point of common observation in saturniids, like several other 
lepidopterans (see Gupta, 1964; Suomalainen, 1979), that the chro- 
mosomes of high-numbered species are always smaller than the chro- 
mosomes of low-numbered species. This evidence and the presence 
of the non-localized nature of centromere, at least at metaphase I, 
probably indicates the occurrence of fission. The possibility of the 
other structural rearrangements like inversions, translocations, etc. 
cannot, however, be disregarded. 


ACKNOWLEDGMENTS 


The authors wish to thank Prof. Dr. S. D. Misra, Head of the Department, for pro- 
viding research facilities and to Mr. S. K. Sarkar, Naturalist, Meghalaya, for providing 
the cocoons of the species. Thanks are also due to Dr. G. S. Arora, Superintending 
Zoologist, ZSI, Calcutta, for helpful information on the systematics. The award of teach- 
er fellowship by the University Grants Commission to one of the authors (RCN) is also 
thankfully acknowledged. 


VOLUME 36, NUMBER 2 BY, 


LITERATURE CITED 


BAUER, H. 1943. Chromosomenforschung. Fortschr. Zool. 7:256—-287. 

BIGGER, T. R. L. 1975. Karyotypes of some Lepidoptera chromosomes and changes 
in their holokinetic organization as revealed by new cytological techniques. Cy- 
tologia 40(3-4):713-726. 

1976. Karyotypes of three species of Lepidoptera including an investigation 
of B-chromosomes in Pieris. Cytologia 41(2):261-282. 

BYTINSKI-SALZ, H. 1938. Untersuchungen an Lepidopteren-hybriden V. Die Ver- 
wandfschaft der Platysamia Arten (Lepidoptera, Saturniidae) nach untersuchun- 
gen uber die Fertilitat und die chromosomen Verhatnisse ihrer Bastarde. Arch. 
Exp. Zellforsch. 22:217-237. 

Cook, M. H. 1910. Spermatogenesis in Lepidoptera. Proc. Acad. Nat. Sci., Philadel- 
phia 62:294—327. 

DANILOVA, L. V. 1973. An electron microscope study of meiosis in diploid males of 
the silkworm. Ontogenez 4:40-48. 

DEDERER, P.H. 1907. Spermatogenesis in Philosamia cynthia. Biol. Bull. 13:94-106. 

1915. Oogenesis in Philosamia cynthia. J. Morpho. 26(1):1-42. 

DEODIKAR, G. B. & C. V. THAKAR. 1958. Cytogenetic studies in Indian silkworms. 
I. Preliminary observations on spermatogenesis in castor silkworm, Attacus ricini. 
Curr. Sci. 27:457-458. 

DEODIKAR, G. B., S. N. CHowpDHuRY, B. N. BHUYAN & K. K. KSHIRSAGAR. 1962. 
Cytogenetic studies in Indian silkworms. II. Chromosome number in muga silk- 
worm, Antherarca assamensis Westwood. Curr. Sci. 31:247-248. 

DEODIKAR, G. B., K. K. KSHIRSAGAR & I. A. KAMATE. 1969. Chromosome number in 
Actias selene Htb—a wild silkworm with reelable cocoons. Ind. J. Genet. Plant 
Breeding 29:126—130. 

ENNIS, T. J. 1976. Sex chromatin and chromosome numbers in Lepidoptera. Can. J. 
Genet. Cytol. 18:119-130. 

GASSNER, G. & D. J. KLEMETSON. 1974. A transmission electron microscope exami- 
nation of hemipteran and lepidopteran gonial centromeres. Can. J. Genet. Cytol. 
16:457-464. 

GupTA, Y. 1964. Chromosomal studies in some Indian Lepidoptera. Chromosoma 
(Berl.) 15:540-561. 

Gupta, M. L. & R. C. NARANG. 1980a. Chromosome number, sex chromatin and sex 
chromosome mechanism in some saturniid moths of India. Entomon 5(1):13-18. 

1980b. Chromosome number and meiotic mechanism of Eriogyna pyretorum 

Westwood (Lepidoptera: Saturniidae). Nat. Acad. Sci. Letters 3(9):279-280. 

1981. Karyotype and meiotic mechanism in muga silkmoths, Antheraea comp- 
ta Roth. and A. assamensis (Helf.) (Lepidoptera: Saturniidae). Genetica 57 (1):21-27. 

JoL.y, M. S., S. K. SEN & S. S. SINHA. 1970. Chromosome number in oak feeding 
tasar silkworm, Antheraea roylei Mr. Curr. Sci. 39:423-424. 

JoLLy, M. S., S. K. SEN, V. SAHAI & G. K. PRASAD. 1977. Chromosome number in 
Antheraea frithi Mr. (Lepidoptera : Saturniidae). Curr. Sci. 46(17):613. 

1978. Chromosome number of Antheraea sivalica Mr. (Lepidoptera: Satur- 
niidae). Sci. and Cult. 44(7):317-318. 

KAWAGUCHI, E. 1933. Die heteropyknose der Geschlechts-Chromosomen bei Lepi- 
dopteren. Cytologia 4:339-354. 

1934. Zytologische untersuchungen am seidenspinner und seiner verwandten. 
II. Spermatogenese bei Antheraea yamamai Guerin, A. pernyi Guerin, und ihre 
bastard. Jap. J. Genet. 10:135-151. 

KERNEWITZ, B. 1915. Spermeogenese bei Lepidoptera mit besonderer Beruchsi- 
chtigung der chromosome. Arch. Naturgeschichte, A, 81:1-34. 

MAKINO, S. 1951. An Atlas of the Chromosome Numbers in Animals. Iowa State 
College Press. 

NARANG, R. C. & M. L. Gupra. 1979a. Chromosome number of Cricula trifenestrata 
Helf. (Lepidoptera: Saturniidae). Curr. Sci. 48(10):465—-466. 


118 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


1979b. Chromosome number of a wild silkmoth, Dictyoploca cachara Mr. 

(Saturniidae, Lepidoptera). Nat. Acad. Sci. Letters 2(5):201. 

1979c. Note on the chromosomes of a wild silkmoth, Loepa katinka. Ww. 

(Lepidoptera: Saturniidae). Chromo. Inf. Serv. 26:17-18. 

1979d. Chromosome studies including a report of B-chromosome in a wild 
silkmoth, Sonthonnaxia maenas (Doubleday) (Saturniidae: saturniinae). J. Res. 
Lepid. 18(3):208-211. 

PARISER, K. 1927. Die cytologie und Morphologie der Triploiden Intersexe des rtick- 
gekreuzten Bastards von Saturnia pavonia L. und S. pyri Schiff. Z. Zellforsch. 
53:415-447. 

RisHi, S. & K. K. RisHi. 1979. Chromosomal analysis of Trabala vishnu Lef. (Lasio- 
campidae, Lepidoptera) with clear indications of localized centromeres. Cytobios 
24(93):33-42. 

SINHA, S. S. & M.S. JOLLY. 1967. Chromosome number in tasar silkworm, Antheraea 
mylitta Drury. Curr. Sci. 36:359. 

SUOMALAINEN, EF. 1969. Chromosome evolution in the Lepidoptera. Chromosomes 
Today 2:131-138. 

TEMPLADO, J., J. ALVAREZ & E. ORTIZ. 1975. Biological and cytogenetical observa- 
tions on Graellsia isabelae (Graellis, 1849) (Lepidoptera: Saturniidae). Eos. Rev. 
Exp. Entomol. 49(1-4):285-292. 

White, M. J. D. 1957. Some general problems of chromosomal evolution and spe- 
ciation in animals. Surv. Biol. Progr. 3: 109-147. 

1973. Animal Cytology and Evolution. 3rd ed., Cambridge Univ. Press, Cam- 

bridge. 961 pp. 


Journal of the Lepidopterists’ Society 
36(2), 1982, 119-120 


ADDENDUM AND CORRIGENDA TO “CLASSIFICATION OF 
THE SUPERFAMILY SESIOIDEA” 


JOHN B. HEPPNER AND W. D. DUCKWORTH 
Department of Entomology, Smithsonian Institution, Washington, D.C. 


ABSTRACT. The subgenera Dipchasphecia, Ductispina, Paradipsosphecia, Pseu- 
dosphecia, and Synansphecia are synonymized with Bembecia in Sesiidae. The names 
Bembecia reisseri Capuse, Chamaesphecia rangnovi Capuse, Homogyna porphyractis 
Meyrick, Microsynanthedon tanala Minet, and Sesia okinawana (Matsumura) are added. 
A number of other changes and corrections are noted. 


In a recently completed paper on Sesioidea (Heppner & Duck- 
worth, 1981), a major reference to Sesiidae (Capuse, 1973) was inad- 
vertantly left out of the world review for the family Sesiidae. Thus, 
we wish to make note of the following taxa as an addition to our 
review of the superfamily. 

The following generic names, proposed as subgenera, are hereby 
placed as synonyms of Bembecia: 


Pseudosphecia Capuse, 1973b:147, new synonymy 
Type-species: Dipsosphecia tenebrosa Pungeler, 1915, by original 
designation [described as a subgenus of Bembecia]. 

Ductispina Capuse, 1973b:149, new synonymy 
Type-species: Dipsosphecia turcmena Bartel, 1912, by original des- 
ignation [described as a subgenus of Bembecia]. 

Paradipsosphecia Capuse, 1973b:150, new synonymy 
Type-species: Dipsosphecia barbara Bartel, 1912, by original des- 
ignation [described as a subgenus of Bembecia]. 

Dipchasphecia Capuse, 1973b:161, new synonymy 
Type-species: Dipsosphecia roseiventris Bartel, 1912, by original 
designation [described as a subgenus of Chamaesphecia; type- 
species now placed in Bembecia]. 

Synansphecia Capuse, 1973b:166, new synonymy 
Type-species: Sesia triannuliformis Freyer, 1845, by original des- 
ignation [described as a subgenus of Chamaesphecia; type-species 
now placed in Bembecia|. 


The following additions and corrections are also noted: 


9. Change setae D2 to D1 and D1 to D2 at the end of the larval diagnosis. 

15. Change Phycodes eucallynta Meyrick, 1937, to Meyrick, 1937a. 

Under current taxa, change 1063 to 1066 species and change 83 to 84 species. 
20. Under current taxa, change 782 to 786 species. Under Bembecia, change 58 to 59. 
21. Under Homogyna, change 9 to 10; under Microsynanthedon, change | to 2. 
27. Under Sesia, add Spherodoptera [sic] Matsumura, 1931a:1017, [misspelling]. 


Sov yU 
— 
(ep) 


120 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


p. 28. Add Sesia okinawana (Matsumura, 1931a:1017), Spherodoptera [sic], [mew 
combination], as a valid species from the Oriental region. Change Sesia montelli 
(Lofquist, 1922:82), Aegeria from a valid species to a junior synonym of Sesia 
bembeciformis (Hubner). 

p. 30. Change Synanthedon danieli Capuse, 1973, to Capuse, 1973a. 

p. 35. Under Chamaesphecia balcanica Zukowsky, 1929, add C. rangnovi Capuse, 
1973b: 167 [nomen nudum], as a synonym. Change C. corsica (Staudinger, 1856) 
to a synonym of C. meriaeformis (Boisduval, 1840), since the latter is not a 
nomen nudum, due to a short description in a footnote. Chamaesphecia mer- 
iaeformis becomes the valid name for the species. 

p. 36. Change Chamaesphecia monspeliensis (Staudinger, 1856) to synonymy with 
C. tengyraeformis (Boisduval, 1840), since the latter is not a nomen nudum, 
due to a short description in a footnote. Chamaesphecia tengyraeformis be- 
comes the valid name for this species. 

p. 37. Change Scalarignathia Capuse, 1973, and Scalarignathia kaszabi Capuse, 1973, 
to Capuse, 1973a. 

p. 39. Add Bembecia reisseri (Capuse, 1973b:156), Pyropteron [new combination], 
as a valid species from the Palearctic region. 

Under B. chrysidiformis (Esper, 1782), B. polistiformis (Boisduval, 1840) is 
not a nomen nudum due to a short description as a footnote in the original 
article. 

Change Bembecia bestianaeli Capuse, 1973, Bembecia dancaudani Capuse, 
1973, Bembecia hannemanni Capuse, 1973, and Bembecia ili Capuse, 1973, in 
each case to “(Capuse, 1973a), Dipsosphecia [new combination].” 

p. 40. Since Chamaesphecia tengyraeformis (Boisduval) 1840 (Sesia) is now a valid 
name and not a nomen nudum, Bembecia tengyraeformis (Herrich-Schaffer, 
1851) (Sesia) becomes a junior homonym, to be replaced by Bembecia san- 
guinolenta (Lederer, 1853). The latter name becomes the valid name for this 
species. 

p. 42. Add Microsynanthedon tanala Minet, 1976:40, for the Ethiopian region. 

p. 43. Add Homogyna porphyractis IMlesaielle ISMN, as a valid species from the 
Ethiopian region. 

p. 49. Change Brenthia dendronympha Meyrick, 1937, to Meyaee 1937a. 

p. 50. Change Brenthia spintheristis [sic] Meyrick to B. spintheritis. 

p. 51. Under Millieria dolosana (Herrich-Schaffer) add M. dolosalis (Heydenreich, 
1851:63), Choreutis [nomen nudum], as another synonym. 

p. 52. Change Tebenna chrysotacta [sic] (Meyrick) to T. chrysostacta. 

p. 62. Add the Capuse reference noted in this paper under the other Capuse, 1973a, 
reference and indicate it as 1973b. 

p. 75. Add the following reference under Meyrick and change the listed 1937 refer- 
ence to 1937a: 1937b. Aegeriadae. Exotic Microlepidoptera 5:119. 

Add the Minet reference as listed below under literature cited. 


ACKNOWLEDGMENT 


We thank P. Leraut, Saint-Maur, France, for noting some of the errors corrected 
above. 


LITERATURE CITED 


CAPUSE, I. 1973. Zur Systematik und Morphologie der Typen der Sesiidae (Lepi- 
doptera) in der R. Pungeler-Sammlung des Zoologischen Museums zu Berlin. Mitt. 
Munchner Ent. Ges. 63:134-171. 

HEPPNER, J. B. & W. D. DucKworTH. 1981. Classification of the superfamily Se- 
sioidea (Lepidoptera: Ditrysia). Smithsonian Contr. Zool. 314: 1-144. 

MINET, J. 1976. Contribution a |’étude des microlépidoptéres de Madagascar. Bull. 
Soc. Ent. France 81:40-43. 


Journal of the Lepidopterists’ Society 
36(2), 1982, 121-131 


EXPERIMENTAL HYBRIDIZATION BETWEEN PHYCIODES 
THAROS AND P. PHAON (NYMPHALIDAE) 


CHARLES G. OLIVER 
R. D. 1, Box 78, Scottdale, Pennsylvania 15683 


ABSTRACT. Phyciodes tharos and P. phaon are common species that occur sym- 
patrically over much of the southern United States. The species differ in larval, pupal, 
and adult phenotypic appearance, habitat preference, and larval foodplant. F, hybrids 
and backcrosses between the species showed an unusual pattern of incompatibility, 
with relatively slight hybrid breakdown in one direction of the cross and total invia- 
bility in the reciprocal cross. The results differ strongly from those obtained in crosses 
between P. tharos and other Phyciodes species. 


In the southern part of the southestern United States Phyciodes 
tharos Drury and P. phaon Edwards are the dominant Phyciodes 
species. They are closely related taxonomically, but while P. tharos 
ranges northward into southern Canada, P. phaon is confined to the 
South and Southwest and ranges into Central America. Except for P. 
phaon, all of the half-dozen or so members of the P. tharos species 
group feed on asters in the larval stage. The larval foodplant of P. 
phaon is Lippia (Verbenaceae), an herbaceous perennial found com- 
monly along sandy roadsides in the Deep South. 

In northern Florida (e.g., Alachua, Bradford, and Levy Cos.) P. thar- 
os and P. phaon are often sympatric along roadsides and in other open, 
sandy areas, but while P. phaon seems fairly closely restricted to this 
habitat, P. tharos is common also in moist grassy fields, lawns, and 
pine-palmetto savannah. Laboratory experiments indicate that P. thar- 
os larvae feed readily on a wide array of Aster species, and that each 
habitat contains suitable foodplants. 

Both species are multivoltine, with the spring emergence begin- 
ning in mid-March in northern Florida. First generation adults of both 
species are of the extreme “spring” phenotype (“marcia’ in P. tharos, 
“hiemalis” in P. phaon) with much more extensive dark markings on 
the ventral hindwing than in the “summer” phenotype (“morpheus’”’ 
and “phaon,”’ respectively). Although fairly similar in appearance, 
adults of the two species are easily distinguished by differences in 
color pattern (Fig. 1, Table 1). The larvae also differ in appearance 
(Table 1), although in general that of P. phaon is more similar in 
appearance to P. tharos than are those of P. campestris Behr (Oliver, 
1978) or P. batesii Reakirt (Oliver, 1979a). In addition first instar lar- 
vae of both P. campestris and P. batesii form rudimentary communal 
webs, whereas P. tharos and P. phaon do not. The haploid chromo- 
some numbers of both latter species is 31 (Maeki and Remington, 
1960). There are apparently no records of natural hybrids. The rela- 


122 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Fic. 1. Parental-type, F, hybrid, and backcross adults: Row A, P. phaon; B, P. 
tharos; C, F, hybrids P. phaon 2 x P. tharos 6; D, backcrosses (P. phaon 2 x P. tharos 
3) 2 xP. tharos 3. Specimens show, left to right: male dorsal, female dorsal, male 
ventral, female ventral. 


tively close taxonomic relationship of P. tharos and P. phaon, together 
with the unusual foodplant of P. phaon, made it seem especially de- 
sirable to me to investigate the relationship between these two species 
as part of my ongoing study of the evolutionary genetics of the P. 
tharos group. 


METHODS AND MATERIALS 


Laboratory stock was derived from one female of P. tharos taken 
17 March 1979 in Gainesville, Alachua Co., Florida; two females of 
P. tharos and one male and two females of P. phaon taken 16 and 21 
March 1979 four mi. west of Otter Creek, Levy Co., Florida; and from 
one male and four females of P. tharos and three females of P. phaon 
taken 17, 18, and 24 March 1979 three mi. north of Waldo, Bradford 
Co., Florida. Cultures at the University of Florida were maintained 
at 25°C and under approximately natural photoperiod conditions (for 
April at latitude 29°30'N). On 19-21 April 1979 the cultures were 
transferred to my laboratory in Pennsylvania, where they were main- 
tained at 27°C days, 24°C nights, and given 16 h light/24 h using rows 
of fluorescent tubes. 


123 


VOLUME 36, NUMBER 2 


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124 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Matings were made by the hand-pairing method (Clarke, 1952) or 
in small cages. Mated females were housed in 10 X 20 cm glass cyl- 
inders and given potted plants or cut sprigs of various asters or Lippia 
nodiflora (L.) Michx. for oviposition. Eggs were removed daily and 
counted, and the substrate leaf kept fresh until the eggs hatched. 
Larvae of P. tharos were reared on cut sprigs of Aster in 10 x 20 cm 
glass cylinders with screening over the tops; whereas, P. phaon were 
reared on cut sprigs of Lippia in 7 x 10 cm closed plastic boxes, since 
the Lippia plants tended to desiccate if exposed to the open air. After 
transferral of the cultures to Pennsylvania all hybrid larvae were reared 
on Aster because of the unavailability of Lippia. For this same reason 
no control broods of P. phaon were reared in Pennsylvania after early 
May. 

F, progeny of wild-collected wild-mated females or (in two cases) 
unmated, wild-collected females were used for the hybrid pairings 
and as parental-type stock for backcrosses. No stock used was inbred. 
Observations were made on parental population, F, hybrid, and back- 
cross phenotypic appearance and larval foodplant acceptance, inter- 
specific courtship behavior, development times and adult eclosion 
patterns, fertility, adult sex ratios, and embryonic, pupal, and eclosing 
adult viability. Single species controls were reared simultaneously for 
comparison with hybrid and backcross broods. 

Data on egg fertility, viability, and sex ratios were treated statisti- 
cally using the Wilcoxon Two-Sample Test. Adult fertility was mea- 
sured by a count of the number of visibly developing eggs divided 
by the total number of eggs laid after a single mating. Development 
times from hatching of the egg to eclosion of the adult were estimated 
by calculating the 99% confidence intervals for the medians of the 
distributions (Owen, 1962). Distributions of development times with- 
in broods or series of broods have been represented by adult eclosion 
graphs showing the number of adults emerging from pupae each day. 


RESULTS 


Interspecific Courtship Behavior 


Males of both species show vigorous courtship behavior when caged 
with females of their own or of the other species. Females apparently 
rarely accept males of other species, however, and only one interspe- 
cific pairing, P. phaon 2 X P. tharos 3, was made without forced 
pairing. 


Phenotypic Appearance 


Differences in phenotypic appearance of the fifth instar larvae, pu- 
pae, and adults of the parental species and their F, hybrid (P. phaon 


VOLUME 36, NUMBER 2 125 


2 x P. tharos 3) are summarized in Table 1. Adults are shown in Fig. 
1. The backcross (P. phaon 2 X P. tharos 3) 2 x P. tharos 3 gener- 
ally resembled P. tharos except for the following characters: 1) larva— 
width of lateral light stripes slightly wider than on P. tharos; 2) adult— 
color of ventral antennal club dark, sometimes with a light tip; 3) color 
of dorsal and ventral forewing intermediate between P. tharos and P. 
phaon; 4) color of ventral hindwing with ground lighter than P. tharos, 
often with a slight, diffused central tawny spot; discal and ventral 
markings more like P. tharos but with some P. phaon influence pres- 
ent; dark markings more sharply defined than on P. tharos. The larva 
and pupa of P. phaon and the larva of P. tharos have been figured by 
Emmel and Emmel (1973) and the pupa of P. tharos by Holland (1931). 

In nature both P. tharos and P. phaon show seasonal polyphenism 
regulated by photoperiod (Oliver, 1976, unpubl. data). In both species 
the short-day forms, which fly in fall and spring, have the underside 
of the hindwings suffused with dark brown or violet-brown, which 
obscures the other dark markings. The character of this suffusion is 
closely similar in both species. Under the laboratory conditions de- 
scribed above only light-colored long-day forms were present in the 
parental cultures. The males in the F, hybrid broods were only of the 
long-day form; varying proportions of the females were of the short- 
day form (Brood No. 79-6: 23.3%; 79-17: 0%; 79-18: 13.1%; 79-34: 0%; 
79-35: 17.6%; 79-36: 71.1% [see Table 4 for numbers of females in- 
volved]). A few backcross females showed moderate expression of the 
short-day phenotype. 


Foodplant Acceptance 


In laboratory oviposition tests P. phaon accepted only Lippia, P. 
tharos only Aster. F, hybrid females (P. phaon 2 x P. tharos ¢) ac- 
cepted either plant readily. 

Newly hatched larvae without feeding experience given the food- 
plant of the other species fed to a very limited extent and died without 
increase in size. F, hybrid larvae of the cross P. phaon & x P. tharos 
6 fed readily on either foodplant. One early brood of this cross was 
split into Lippia- and Aster-feeding groups. Growth was markedly 
slower on Lippia, and the Lippia-feeding group was changed to Aster 
when the cultures were moved to Pennsylvania. Part of one brood of 
the backcross (P. phaon 2 x P. tharos 3) ° xX P. tharos 3 was started 
on Lippia and grew until the second instar, when all of the larvae 
(N = 40) died. 


Viability, Fertility, and Sex Ratio 


Embryonic viability in the parental control broods was extremely 
high. In the F, hybrid (P. phaon 2 x P. tharos ¢) there was significant 


126 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


reduction in viability (Table 2). In the reciprocal hybrid there was 
either complete very early embryonic inviability or a complete lack 
of egg fertilization following insemination. Females mated in this cross 
oviposited normally (in large egg patches), rather than in the pattern 
demonstrated by uninseminated females (very few eggs laid, widely 
scattered over the foodplant). Since in other Phyciodes crosses this 
has proved to be an infallible indicator of mating success, these mat- 
ings were assumed to be successful even though the presence of sper- 
matophores was not verified. The backcross (P. phaon @ x P. tharos 
3) 2 x P. tharos 3 showed a significant reduction in visible egg fer- 
tility and reduced embryonic viability. The reciprocal backcross P. 
tharos 2 Xx (P. phaon 2 X P. tharos 3) 3 showed massive reductions 
in egg fertility and complete embryonic inviability. The reduction in 
visible egg fertility in these crosses may have been due to very early 
embryonic inviability or to reduced parental fertility. No stock of P. 
phaon could be maintained in Pennsylvania for backcrosses because 
of a lack of Lippia for foodplant. 

Post-larval (i.e. prepupal, pupal, and ecdysing adult) viability was 
significantly lower for the P. phaon parental broods than for P. tharos 
(P = .005) (Table 3). This may have been due to the different larval 
rearing containers for P. phaon, since humidity in these containers 
was very high. F, hybrid and backcross broods were reared in the 
same manner as P. tharos; post-larval viabilities of these two series 
of broods were significantly lower than for P. tharos (P = .005 for both 
values) but not lower than both P. tharos and P. phaon considered 
together. It is possible that neither parental foodplant was ideal for 
the hybrid larvae, and that some reduction in viability was due to this. 

Adult sex ratios in the F, hybrid broods and in the backcross broods 
showed no change from those of the parental control broods (Table 
3). 


Development Times and Eclosion Graphs 


Development times from hatching of the egg to eclosion of the adult 
were about the same for the two parental species, although some 
broods of P. phaon averaged a day or so faster than P. tharos (Table 
4). Development times of the P. phaon broods varied more than did 
those of P. tharos. 

In F, hybrid broods of the cross (P. phaon 2 x P. tharos 3) reared 
on Aster, males showed development times similar to those of the 
control broods, but those of the F, females averaged at least several 
days longer than those of the controls. Development times of both 
males and females of this F, hybrid tended to vary more than those 
of the controls. In addition, the eclosion graphs of both sexes showed 


27 


VOLUME 36, NUMBER 2 


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128 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 4. Development times in days from hatching of egg until eclosion of adult 
for P. tharos and P. phaon control broods, F, hybrid (P. phaon @ x P. tharos 3) and 
backcrosses ({[P. phaon @ X P. tharos 3] 2 x P. tharos 3). “Medians” show 99% con- 
fidence limits. See text for rearing conditions. Mr = March, A = April, My = May, 


= June. 
Males Females 
Brood no. Date hatched N Min—Max Median N Min—Max Median 
P. tharos 
79-3 29 Mr od 57 26 43 24-30 WO PAT 
79-5 29 Mr-3 A 16 25=—29 26—26 18 WAS), 26-28 
79-9 27-29 Mr 111 24-30 26 125 US —o) ON 
79-10 30 Mr-1 A 37 25—26 AS HAS 2th 25-28 25-26 
79-24 4 My 47 18-20 19 30 19-24 19 
79-25 1-4 My 54 18-21 19-21 32 192i 19-20 
79-26 4 My 33 18-19 18 20 18-19 18-19 
P.. phaon 
79-7 29 Mr-1A 30 23-30 24-26 34 25-29 26—271, 
79-12 31 Mr 23 23-24 23 34 YEU) Ue ve! 
79-13 12 A 35 23-26 24 30 JZA—31 24-26 
F, Hybrids 
79-6 29 Mr-1A 44 25—309 29-30 29 OS) 40-43 
719-17 1-6 My 35 19-26 20 65 All 8\L 23-24 
79-18 30 A-5 My 64 17-24 O= Fil 63 21-29 Wo) 
79-34 8-11 My 10 19-24 19-24 7 22-29 22-16 


79-35 7-11 My 56 18-27 19-20 93 20-30 23-26 
79-36 6-10 My 58 18-44 19-22 45 22-33 25-28 


Backcrosses 
79-47 oie) 9 IPO 17-20 16 19-24 19-24 
79-49 4-5 J 50 16-20 17-18 49 17-26 18-19 
79-50 5 | 54 15-20 eas 83 Wi 22 19-20 
79-51 4-5 J 28 16=23 Ike 23 18-23 19-21 
79-55 7-8 J 67 YS) 19-20 61 18-24 20-21 
79-58 10 J 2D, 15-20 17-19 20 18-21 19-20 


a tailing-off effect. This was more marked in the females (Fig. 2). 
Because of slightly different rearing conditions, broods hatched dur- 
ing March and early April cannot be compared with those hatching 
during May and June. Development times of the backcrosses to P. 


tharos were significantly shorter than those of P. tharos or of the F, 
hybrids. 


DISCUSSION 


At first glance the results seem to give a somewhat contradictory 
picture of incompatibility between Phyciodes tharos and P. phaon. 
In the F, hybrid (P. phaon 2 x P. tharos ¢) and its backcross to P. 
tharos 3, hybrid sex ratios are normal and development times only 
slightly affected; whereas, the reciprocal hybrid is totally inviable, 


VOLUME 36, NUMBER 2 129 


80 


60 


Bas) 
(—} 


Adults Eclosing 


20 10 


30 40 50 


Days until Eclosion 


Fic. 2. Distributions of times required for development of typical P. tharos, P. 
phaon, F, hybrid, and backcross broods from hatching of eggs until eclosion of adults. 
A, P. tharos, Brood 79-9; B, P. phaon, Brood 79-7; C, F,; hybrid P. phaon 2 x P. tharos 
36, Brood 79-6; D, backcross (P. phaon 2 x P. tharos 3) 2 x P. tharos 3, Brood 79-55. 
(See Table 4 and text for rearing conditions and dates.) 


and the backcross P. tharos 2° x (P. phaon 2 x P. tharos 3) 3 almost 
so. It would appear from these results that, while the nuclear materials 
of P. tharos and P. phaon are quite compatible (i.e. can cooperate to 
direct harmonious growth and development) and while P. tharos nu- 
clear material is relatively compatible with P. phaon or hybrid cyto- 
plasm, P. phaon nuclear material is highly incompatible with P. thar- 
os cytoplasm. This incompatibility may involve crucial differences in 
one or more of the many factors that determine the composition of 
the cytoplasmic environment in which foreign nuclear material must 


130 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


function to produce a viable hybrid individual organism. An incom- 
patibility of this sort may be ultimately attributable to a relatively 
slight degree of differentiation in gene regulation (see Oliver, 1979b, 
for a fuller discussion) and does not contradict the other results pre- 
sented here. The conclusion is, then, that P. phaon and P. tharos show 
relatively slight overall genetic differentiation. Hybridization in na- 
ture is probably prevented by barriers involving courtship behavior. 

The pattern of hybrid incompatibility in the present series of cross- 
es is quite different from that shown in hybridization experiments 
using P. tharos and P. campestris (Oliver, 1978), P. batesii (Oliver, 
1979a), or the entity I have referred to as P. “tharos Type B” (Oliver, 
1979a, 1980). These combinations show a homogeneous pattern of 
effects, which involves mainly slight to moderate (but not total) re- 
duction in viability in the F, hybrids and backcrosses, skewed F,, adult 
sex ratios, and abnormal F, hybrid development times. In the F, hy- 
brid females eclosion occurs before that of the males when P. tharos 
is the male parent. In the reciprocal crosses female eclosion is de- 
layed, and male and female curves usually do not overlap. In the (P. 
phaon 2 xP. tharos 3) F, hybrid broods, however, females show 
slightly delayed rather than speeded up eclosion times. These ab- 
normalities in development time probably are in some way related to 
parental species differences in larval diapause induction thresholds 
(Oliver, unpubl. data). | 

Expression of the “marcia’’-“hiemalis” phenotypes in the female 
F, hybrids may indicate that this form results unless the “morpheus ’ - 
“phaon” form is induced. One possible explanation for failure of this 
induction in female hybrids is that the “switch” gene and/or modifiers 
are linked with both a diapause induction-development rate gene 
complex and with sex. Since it is the females that are heterogametic 
in Lepidoptera, in this cross the female “morpheus -“phaon” form 
must be induced in P. phaon cytoplasm using regulation by P. tharos 
genetic material, this induction fails in a high percentage of individ- 
uals. This case seems to be analogous to that in the cricket genus 
Pteronemobius (Masaki, 1978), where the F, hybrid males (the het- 
erogametic sex in Orthoptera) show abnormal growth rates and pho- 
toperiodic responses. 

The pattern of hybrid incompatibility between P. tharos and P. 
phaon differs also from that in butterfly hybrids outside the genus 
(reviewed in Lorkovic, 1978, and Oliver, 1979b). In general these 
latter show incompatibility similar to that in the other Phyciodes 
crosses discussed above. I know of no case in which the reciprocal 
F, hybrids differ so drastically in viability as do those between P. 
tharos and P. phaon, although recent crosses between Pieris callidice 


VOLUME 36, NUMBER 2 131 


Hubner and P. occidentalis Reakirt (Pieridae) (Shapiro, 1980) show a 
basic similarity that may be due to the same genetic effects. 


ACKNOWLEDGMENT 


I am grateful to the Department of Zoology, University of Florida, Gainesville, Flor- 
ida, for the use of laboratory facilities during my visit there in March and April 1979. 


LITERATURE CITED 


EMMEL, T. C. & J. F. EMMEL. 1973. The butterflies of Southern California. Nat. Hist. 
Mus. L. A. Co., Sci. Ser. 26:1-148. 

HOLLAND, W. J. 1931. The Butterfly Book, revised ed. Doubleday & Co., New York. 
424 pp. 

LORKOVIC, Z. 1978. Types of hybrid sterility in diurnal Lepidoptera speciation and 
taxonomy. Acta Entomol. Jugoslavica 14:13-24. 

MAEKI, K. & C. L. REMINGTON. 1960. Chromosomes of North American Lepidoptera. 
Part 4. J. Lepid. Soc. 14:179-201. 

MASAKI, S. 1978. Seasonal and latitudinal adaptations in the life cycles of crickets. 
In Evolution of Insect Migration and Diapause (H. Dingle, ed.). Springer-Verlag, 
New York. 284 pp. 

OLIVER, C. G. 1978. Experimental hybridization between the nymphalid butterflies 
Phyciodes tharos and P. campestris montana. Evolution 32:594-601. 

1979a. Experimental hybridization between Phyciodes tharos and P. batesii 

(Nymphalidae). J. Lepid. Soc. 33:6—20. 

1979b. Genetic differentiation and hybrid viability within and between some 

Lepidoptera species. Amer. Natur. 114:681-694. 

1980. Phenotypic differentiation and hybrid breakdown within Phyciodes 
“tharos” (Lepidoptera: Nymphalidae) in the northeastern United States. Ann. 
Entomol. Soc. Amer, 73:715—721. 

OwEN, D. B. 1962. Handbook of Statistical Tables. Addison-Wesley, Reading, Mass. 
580 pp. 

SHAPIRO, A. M. 1980. Genetic incompatibility between Pieris callidice and Pieris 
occidentalis nelsoni: differentiation within a periglacial relict complex (Lepidop- 
tera: Pieridae). Canad. Entomol. 112:463-468. 


Journal of the Lepidopterists’ Society 
36(2), 1982, 132-135 


NOTES ON THE BIOLOGY OF 
ZEGRIS EUPHEME (PIERIDAE) 


STEVEN COURTNEY 


Department of Zoology, University of Liverpool, P.O. Box 147, 
Brownlow Street Liverpool L69 3BX, ENGLAND 


By comparison with the much researched Pierini, the ecology and 
behavior of Euchloini are rather poorly known. Only for the Palaearc- 
tic Anthocharis cardamines L. have detailed studies been made (Wil- 
liams, 1915; Wiklund & Ahrberg, 1978). In this note I record some 
observations made in Morocco during 1978 on another euchloinid, 
Zegris eupheme Lederer, which are of interest for their similarities 
and contrasts with A. cardamines and with Pierini. 

Z. eupheme is a large handsome insect, of disjunct distribution in 
the Palaearctic, being found in southern Spain and Morocco (subspe- 
cies meridionalis), and southern U.S.S.R., Asia Minor and the Near 
East (subspecies eupheme). It is unusual among the Pierinae in being 
particularly associated with a single crucifer species, Isatis tinctoria 
L., although it may use other Cruciferae as larval host plants (Powell, 
1932). In the Middle Atlas Mountains of Morocco, the butterfly flies 
during late April and May and may often be seen in areas where the 
foodplants are growing. At this time of year the growth of vegetation 
is rich and verdant in valley bottoms; spring is the time of greatest 
floral and faunal abundance (thereafter the high summer temperatures 
give a very desiccated-looking landscape). I. tinctoria, a large, yellow- 
flowered crucifer about 1 m in height, grows in peripheral open areas 
around valleys, including roadsides and stony banks, where vegeta- 
tion is otherwise short. At Ifrane, in the Middle Atlas, Z. eupheme 
has been associated with I. tinctoria since at least 1965, when the 
late Baron de Worms (1965) recorded flourishing populations of but- 
terfly and crucifer in a hollow next to the Ballima Hotel. On subse- 
quent visits he confirmed the continuing success of both populations, 
which were both still abundant in 1978. Most of the observations 
recorded here were made in this small area (about 100 m square); 
however, I. tinctoria occurred in sporadic small groups elsewhere, 
and Z. eupheme was very wide-ranging and could be found in almost 
any habitat in the area (ranging from rocky slopes to cedar forest). 

As to be expected from its size, Z. ewpheme is a very fast flier, 
making it impossible to net by pursuit. Like the satyrid Eumenis se- 
mele (L.), it often evades capture by dropping below the sweep of 
the net. The only successful method of capture is to station oneself 
in areas where Z. eupheme is flying and try ‘head on’ shots. The males 


VOLUME 36, NUMBER 2 133 


appear to ‘patrol’ (sensu Scott, 1974) areas in search of females, and 
during such flight they investigate any flying insects of the approxi- 
mate size and color of Z. eupheme. Thus, males were seen to chase, 
vigorously and persistently, individuals of the white pierids, Pieris 
brassicae L. and Artogeia rapae (L.), and the yellowish pierids, Go- 
nepteryx rhamni L. (both the primrose colored male, and the paler 
female) and Colias crocea Gffy. All such interactions were initiated 
by Z. eupheme, although other pierids are equally investigative in 
other situations. Several interactions were seen between males of Z. 
eupheme, but only one encounter was observed between sexes, which 
happened to produce a successful courtship and copulation. On being 
contacted the female flew about 10 m, with the male spiralling vig- 
orously around her. She then landed, with the male continuing in the 
air and after a few seconds took off again. Once more the male ap- 
peared to force her to the ground but did not settle himself, fluttering 
around her head. This process was repeated several times, and on 
each occasion the pair moved a few meters. Eventually, the male 
settled down in a head to head position and rapidly moved around 
and initiated copulation with the now quiescent female. At this point 
the animals were collected; dissection of the female revealed no sper- 
matophores, confirming that she was unmated. Seven other females 
each contained a single spermatophore, perhaps indicating that Z. 
eupheme leans toward monogamy as does A. cardamines (Courtney, 
1980). At no time did females adopt the ‘mate-refusal’ posture (raising 
the abdomen vertically, while spreading the wings), a typical reaction 
of other Pierinae females. 

The eggs of Z. eupheme, like those of most Euchloini, are placed 
upon the inflorescences of the crucifer host. When laid the egg is 
white, but it soon develops a red color, as in other Holarctic Euchloini 
(e.g. Williams, 1915). By careful examination of the flowerheads of I. 
tinctoria, it is readily possible to find the eggs. During the inclement 
weather of 28 and 29 April 1978 (snow occurred on May 1), the po- 
sition of every Z. eupheme egg on I. tinctoria at the Ballima Hotel 
site was mapped. It was soon apparent that only certain I. tinctoria 
were chosen as oviposition sites. Eggs were concentrated upon un- 
opened buds, and open flowers were never chosen (Table 1). Similar 
behavior in A. cardamines has been hypothesized to be adaptive, 
leading to larval feeding only on young buds and fruit, which repre- 
sent optimal food (Wiklund & Ahrberg, 1978). Additionally, eggs were 
nonrandomly distributed among foodplants, with plants on the edge 
of the clump receiving many more eggs than those in the center. 
However, since edge plants are automatically those at low density, it 
is impossible to decide whether female Z. eupheme prefer to lay eggs 


134 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 1. Z. eupheme eggs upon I. tinctoria plants of different floral ages. 


Plants with — N ; Number of eggs 
10% flower buds open Sui 85 
10-90% flower buds open 233 19 
Flowers fully open PAIL 0 
Ts 104 


x? = 41.6, P< 0.001 at 2 df. 


on plants at low density or prefer edge plants or both. Fig. 1 presents 
the observed deposition of eggs as a density plot. Similar patterns of 
egg deposition are known for A. cardamines (Wiklund & Ahrberg, 


15 


0-5 


0-4 59 10-19 20-29 30-39 40-49 50-59 60-69 


No. of plants within 2m. 


Fic. 1. The effect of I. tinctoria density (expressed as the number of other individ- 
uals within 2 m) upon the Z. eupheme eggload (eggs/plants). A total of 104 eggs on 731 
plants were found. 


VOLUME 36, NUMBER 2 135 


1978) and for the pierids Artogeia rapae (e.g. Kobayashi, 1965), A. 
napi (in prep.), Pontia protodice and P. occidentalis (Shapiro, 1975). 
Various hypotheses have been suggested for the cause of such ‘edge’ 
effects, including choice by females of areas where parasite attacks 
are rare (Shapiro, 1975) and the incidental effects of female movement 
patterns and searching behavior (Jones, 1977). In such a large, mobile 
butterfly as Z. eupheme, Jones's hypothesis predicts an extremely pro- 
nounced edge-effect, as appears to be the case at Ifrane (a fuller dis- 
cussion of edge-effects in Pierinae, particularly A. cardamines, is in 
preparation). The age of I. tinctoria inflorescences was not related to 
plant density. 

Larvae of Z. eupheme were not studied. Beautifully camouflaged 
crab-spiders, common on the flowerheads of I. tinctoria, were poten- 
tial predators. Powell (1932) records a very high rate of parasitism by 
braconids (Apanteles spp.). 

Z. eupheme adults were only seen to feed from I. tinctoria blooms, 
despite a profusion of other nectar sources available. The adults were 
also frequently found roosting upon the host plant, at which times the 
yellow-green mottling of the hind-wing underside provided admira- 
ble camouflage, blending with the yellow inflorescences. It may be 
that restriction of larval feeding to a single plant species (or predom- 
inantly so) has allowed similar specializations (to predictable re- 
sources) in adult behavior. 


ACKNOWLEDGMENTS 


Although the Ballima Hotel site is openly accessible, I soon became aware that it is, 
in fact, a restricted area, belonging to the Moroccan Army. To the senior officers, who 
accepted my explanations as to cryptic maps (of flowers and butterfly eggs) and released 
both me and my data, I am extremely grateful. This work was carried out during the 
tenure of an S.R.C. Research Studentship. 


LITERATURE CITED 


COURTNEY, S. P. 1980. Studies on the biology of the butterflies Anthocharis carda- 
mines (L.) and Pieris napi (L.) in relation to speciation in Pierinae. Ph.D. Thesis, 
April 1980, Durham University. 

JONES, R. 1977. Movement patterns and egg distributions in cabbage butterflies. J. 
Anim. Ecol. 46:195-212. 

KOBAYASHI, I. 1965. Influence of parental density on the distribution pattern of eggs 
in the common cabbage butterfly. Res. Popul. Ecol. 7: 109-117. 

POWELL, H. 1932. Pupation of Zegris eupheme meridionalis, Led. Proc. Entomol. 
Soc. 6:52—54. 

SHAPIRO, A. M. 1975. Ecological and behavioural aspects of co-existence in six cru- 
cifer-feeding butterflies. Am. Midl. Nat. 93:424-433. 

WIKLUND, C. & C. AHRBERG. 1978. Hostplants, nectar source plants, and habitat 
selection of males and females of Anthocharis cardamines. Oikos 31:169-183. 
WILLIAMS, H. B. 1915. Notes on the life history and variation of Euchloe cardamines 

L. Trans. S. Lond. Nat. Hist. Soc. 62-84. 
WoRMS, BARON DE. 1965. Spring Collecting in Morocco Ent. Rec. 77:177-182. 


Journal of the Lepidopterists’ Society 
36(2), 1982, 136-144 


HOSTPLANT RECORDS AND DESCRIPTIONS OF JUVENILE 
STAGES FOR TWO RARE SPECIES OF 
EUEIDES (NYMPHALIDAE) 


JAMES L. B. MALLET AND JOHN T. LONGINO 
Department of Zoology, University of Texas, Austin, Texas 78712 


ABSTRACT. New hostplant records are presented for two species of Eueides. E. 
lineata feeds on an as yet undescribed species of Passiflora in southern Mexico, and 
E. vibilia feeds on old leaves of Passiflora pittieri on the Osa Peninsula of Costa Rica. 
Behavioral observations are given and juvenile stages described for the two Eueides 
species. Larval mimicry is suggested for E. vibilia and two sympatric species of Hel- 
iconius, and an evolutionary mechanism for the mimicry is discussed. 


Heliconiine butterflies are the best known group of Neotropical 
Lepidoptera. Their taxonomy has been unusually well worked out 
(Emsley, 1963, 1964, 1965; Brown & Mielke 1972; Brown 1976, 1979). 
This work has provided a firm basis for biological study on the species 
(reviewed by Brown, 1981). One of the most important features of the 
work on heliconiines has been their hostplant specialization (Gilbert, 
1975; Benson et al., 1976; Benson, 1978; Smiley, 1978). All known 
species of Heliconiini use hostplants in the closely related families 
Passifloraceae (Benson et al., 1976) or Turneraceae (D. H. Janzen, 
pers. comm.), and are usually specialized within particular subgenera 
of the Passifloraceae (Benson et al., 1976). Relationships between Hel- 
iconius and Passiflora have proved to be useful tools in the taxonomy 
of both groups. We here describe the hostplants and larval stages of 
two little known species of Eueides (Nymphalidae: Nymphalinae: 
Heliconiini): E. lineata and E. vibilia, respectively. The young stages 
of Eueides lineata Salvin & Godman and its hostplants were previ- 
ously unknown. Although the young stages of Eueides vibilia Godart 
are known (Brown, 1981) they have not been described in detail, and 
there are no hostplant records north or west of Guyana (Benson et al., 
1976). We have used a method for description of young stages which 
allows direct comparison with the descriptions of Beebe et al. (1960). 


Localities 


In Mexico we found E. lineata at Playa Escondida near Catemaco, Veracruz (18°30'N, 
95°0’W) and at Laguna Encantada above San Andres Tuxtla, Veracruz (18°30’N, 95°10’W). 
Playa Escondida is on the Gulf coast, and we made our observations at altitudes of 
between 50 and 100 m above sea level where there is some forest and recently cut 
pasture. At Laguna Encantada there are the scrubby remains of forest around the La- 
guna, which now acts as a rather eroded watering hole for cattle; the altitude was 
approx. 600 m. 

The larvae and adults of Eueides vibilia were found at San Pedrillo, Parque Nacional 
Corcovado, Costa Rica (8°38'N, 83°44’W). They were not found at Sirena, which is also 
within the park (8°28’N, 83°35’W). Both of the Corcovado sites are on the Pacific coast 


VOLUME 36, NUMBER 2 Se 


at sea level. All four of the sites have tropical lowland rainforest, but of a type with a 
moderately pronounced dry season. 


Eueides lineata 
A. Distribution and Mimicry 


E. lineata is an exclusively Central American and Mexican butter- 
fly. It is usually found at altitudes below 1000 m (Brown, 1979) and 
participates in the common “orange’ mimicry ring in this area, which 
includes Dryas julia (Fab.), Eueides aliphera (Godart), Eueides lybia 
(Fab.), Eueides vibilia, and Dione juno (Cramer). E. lineata is illus- 
trated in color, together with its co-mimics, in Lewis (1974, pls. 43-44) 
and Smart (1976, pl. 87). The adult Eueides that have been tested are 
distasteful to birds, although they are not rejected as much as Heli- 
conius (Brower et al., 1963); so, the mimicry involved here is probably 
Mullerian. 


B. The Hostplant 


The hostplant of E. lineata at Playa Escondida and Laguna Encan- 
tada is a new species of Passiflora. It was found in 1978 by L. E. 
Gilbert, and he is in the process of describing it. In the remainder of 
the paper it will be referred to as Passiflora, sp. nov. It is an aberrant 
species which has a sticky puberulence on both the leaf surfaces. The 
species produces flowers from the tendrils, which is a characteristic 
of old world genera of Passifloraceae, such as Adenia, and some mem- 
bers of the new world subgenus of Passiflora, Astrophea. Most Pas- 
siflora produce flowers directly from leaf axils. There is a pronounced 
difference in the juvenile and mature vegetation: juvenile leaves have 
filiform petiolar nectaries and variegated leaves; leaves on older plants 
are unvariegated, less puberulent, and have large saucer-shaped pet- 
iolar nectaries. The vine can grow into the canopy of tall trees, but 
we were able to search only small plants in newly felled pastures. 
Adults seemed rare, but a concentration of adults occurred around a 
single canopy-level liana in a tree on the edge of a pasture. On 13 
March 1980 a larva and pupal skin were discovered on the plant at 
Laguna Encantada. On 14 March we carefully searched leaves of young 
plants of Passiflora, sp. nov. at Playa Escondida and discovered one 
third-instar larva, three first-instar larvae, and three eggs of E. lineata. 
We also searched P. serratifolia Linn., common at the site, and found 
many eggs and larvae of Eueides isabella (Cramer) (young stages sim- 
ilar to those described by Beebe et al., 1960), but we found neither 
E. isabella on Passiflora, sp. nov. nor E. lineata on P. serratifolia. L. 
E.. Gilbert (pers. comm.) has reared E. lineata at Monteverde (1300 
m), Costa Rica, on an undetermined Passiflora. The association of E. 


138 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Fic. 1. Adults, top to bottom: Eueides aliphera, E. lineata, E. vibilia (male), E. 
vibilia (female). 


VOLUME 36, NUMBER 2 139 


lineata with such an atypical Passiflora in Mexico makes the discov- 
ery of E. lineata’s hostplants in lower Central America an exciting 
prospect. 


C. The Egg 


The egg of E. lineata is laid singly on the undersides of fully ex- 
panded, full-size leaves of the hostplant, a behavior common to many 
Eueides. The egg is greenish white, dome-shaped, and measures 
0.83—0.84 mm high by 0.77-0.83 mm wide. It is similar in general 
shape to that of other Eueides spp. (Beebe et al., 1960), having in- 
dentations caused by the nurse cells during oogenesis and ridges be- 
tween them. There are 10-11 horizontal ridges in total and 19-20 
vertical ridges (n = 3). 


D. The Larva 


Larvae were reared in the laboratory on cut leaves (fresh daily) after 
being collected at Playa Escondida. Detailed measurements were 
made of fifth-instar larvae and pupae. Larval periods were measured 
(sample sizes in parentheses): egg to pupa, 21-22 days (n = 3); first- 
instar larva to pupa, 18-19 days (n = 3); third instar to pupa, 13 days 
(n = 1); fifth instar to pupa, 6 days (n = 1). The prepupal period lasts 
one day (n = 2), during which time the larva constructs a pad, changes 
to a pale yellow color, and hangs from the pad. 


The fifth-instar larva (n = 1) (Fig. 2) has a maximum length of 24 mm. The head is 
2.5 mm high and of the same width, is colored orange with black spots and white 
markings. The posterior dorsal and lateral border of the head capsule is black. A pair 
of black backwardly curved scoli, 3 mm in length, top the head. Dorsally, segments T1 
to A7 are black and white in transverse stripes, four to a segment, A8 is dorsally orange, 
AQ and A10 are again black and white. Laterally there is a creamy-yellow line on T3—A8 
that includes the spiracles. The underside is transparent greenish yellow, except the 
tips of the prothoracic legs, which are black. Dorsal scoli are black, paler medially; 
T2 = 2.5 mm, T3 = 2.75 mm, Al-A7 = 3.00 mm, A8—A9 = 2.75 mm. Lateral scoli are 
black, T2-T3 = 2.0 mm. Supralateral scoli are black, paler medially; Al = 2.0 mm, 
A2-A5 = 3.0 mm, A6—-A7 = 2.75 mm, A8 = 2.5 mm. Sublateral scoli are pale-translu- 
cent; Al = 1.25 mm, A2 = 2.0 mm, A3—-A6 = 2.25 mm, A7 = 2.0 mm, A8 = 1.75 mm. 
Anal scoli are black, 2.25 mm. The prothoracic plate is in the form of two hourglass- 
shaped plates placed transversely, one each side of the midline. 


In the hanging prepupal stage, all body and head color is lost, the 
larva becoming pale greenish yellow. The black scolus color remains 
with black transversely oblong bases of the dorsal and supralateral 
abdominal scoli, except on A8 (scolus bases are here orange in the 
mature larva). The black head markings also remain. 


140 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Fic. 2. Juvenile stages, clockwise from upper left: Eueides lineata (fifth-instar larva), 
E. vibilia (fifth-instar larva), E. vibilia (pupa), E. lineata (pupa). 


E. The Pupa 
(Fig. 2) 


The pupal period seems variable in this species. We recorded pe- 
riods of 9, 8, 8, 8, 7, 6, 6, and 5 days. In general outline the pupa of 
E. lineata is typical for Eueides, being bent ventrally so that it hangs 
horizontally. 


The cephalic projections are 2.0 mm long, about the same as the diameter of the eye, 
without projections or scallops, smooth compressed laterally, curved dorsally and pos- 
teriorly, distally tapered and pointed. Antennae are without spines. Gold spots are 
absent. T2 has a median crest. The paired dorsal projections on T1 are about 0.5 mm 
long, on T2 nearly absent, and on T3 about 0.25 mm. There are no postmedian tubercles 
on the forewing. The three submarginal tubercles are well developed, the posterior 
two connected to the wing margin by a stripe of black pigment. The veins of the pupa 
are not marked with pigment. The profile of the antennae and wings projects little 
ventrally. The abdomen has paired dorsal projections as follows: Al = 0.25 mm, A2 = 0.5 
mm, A3-A7 = 1.0 mm. Tubercles on A3—A7 are directed anteriorly; evidence of bifidy 
is limited to a small bump on the posterior part of the stem. The general color is white 
marked with brown and olive green, especially a dorsal brown spot on A2—A3, and 
brown-marked scoli on the abdomen. There are some details in black, especially around 
the cremaster. The silk pad color is white. The total length is 15.5-17.0 mm (N = 5). 
The pupa differs from other Eueides in having dorsal paired tubercles A3—A7 short, 
homogeneous, anteriorly directed, and not darkly pigmented. 


Eueides vibilia 
A. Distribution and Adult Mimicry 


In Central America the sexually dimorphic E. vibilia occurs as far 
North as the Sierra de Tuxtla in Mexico (Ross, 1967) and has the 


VOLUME 36, NUMBER 2 14] 


subspecies name vialis Stichel. The orange and black male might 
loosely be described as a mimic of the “orange” mimicry group (see 
above under E. lineata). The female is yellower on the forewing me- 
dial region and has black ray markings on the hindwing, resembling 
Actinote anteas Doubleday (Acraeinae), with which it is sympatric in 
some areas of Costa Rica (P. DeVries, pers. comm.). We did not, how- 
ever, find A. anteas in Corcovado. E. vibilia is a widespread species 
and is found south to Rio de Janeiro, Brazil, where E. v. vibilia fe- 
males, E. pavana Menetries, and Actinote spp. participate in a mim- 
icry ring (Brown and Mielke, 1972). The species is rare in Costa Rica, 
only being known from a handful of specimens. These specimens are 
all from lowland localities on both the Pacific and Atlantic drainages 
of Costa Rica. 


B. The Hostplants 


The three reported hostplants of E. vibilia in South America are 
Mitostemma glaziovii Mast., Passiflora (Astrophea) costata Mast., and 
P. (A.) mansii (Mart.) (Benson et al., 1976). In Costa Rica we discov- 
ered larvae on old leaves of another Astrophea species, Passiflora 
pittieri Mast., while searching at San Pedrillo for larvae of Heliconius 
hewitsoni Staudinger on 13 June 1980. P. pittieri is quite common at 
San Pedrillo and Sirena. 

Although Sirena is only 24 km down the coast from San Pedrillo, 
we have found neither larvae nor the distinctive old-leaf skeletonizing 
damage of E. vibilia at Sirena. There is evidence that P. pittieri has 
a more or less continuous distribution between the two sites. This 
indicates great patchiness of E. vibilia, which clearly does not result 
from its hostplant distribution. P. pittieri, judging from the number 
of herbarium specimens available, is as rare as E. vibilia. P. pittieri’s 
rareness is an artifact of its growth form and reproduction (Longino, 
unpub. data). It flowers rarely, usually in the forest canopy, and it is 
vegetatively very cryptic, juveniles looking more like understory tree 
seedlings than Passiflora. E. vibilia’s rareness is no doubt real, since 
the Osa Peninsula has been visited for many years by lepidopterists 
and students of heliconiine biology (L. E. Gilbert, W. W. Benson, P. 
DeVries, J. Smiley), and this is the first report of E. vibilia on the 
Osa. 


C. The Egg 


A batch of 74 eggs is figured by Benson et al. (1976) which was laid 
on a mature Passiflora mansii leaf underside in Mato Grosso, Brazil. 
We have not observed oviposition, but assume from the gregarious 
larvae that E. vibilia lays eggs in batches in Costa Rica also. According 


142 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


to Brown (1981) the eggs are red and white, measure 1.0 mm high by 
0.8 mm wide, and have 14 vertical and 8 horizontal ridges. 


D. The Larva 


Larvae were reared on cut leaves after collection at San Padrillo. 
The larval periods were: third instar, 3 days, fourth instar, 3 days 
(n = 1 batch), fifth instar, 5-7 days (n = 2 batches). First-instar larvae 
were not observed. Second to fourth instar had shiny black head cap- 
sules and black head scoli, with pale yellowish-green bodies and pale 
body scoli. In this way they resembled the larvae of the sympatric H. 
hewitsoni, that also feed gregariously on P. pittieri in the same hab- 
itat. The young larvae of E. vibilia can be distinguished from those 
of H. hewitsoni by the larger head scoli of the former and their habit 
of skeletonizing older leaves, rather than eating young shoots. This is 
probably an example of larval mimicry. During the fourth instar the 
scoli and upper body surface begin to darken, and the fifth instar is 
non-mimetic. 


The fifth instar (n = 1) (Fig. 2) extends to about 23 mm long. The head is 3 mm high 
and broad, is black and has a pair of black dorsal scoli about 4 mm in length. Dorsally, 
the larva is olive green with black patches. Laterally, there is a creamy-yellow line that 
includes the spiracles; ventrally, the color is translucent greenish yellow. The prolegs 
are: Tl = black, T2 = translucent with black tips, T3 = translucent. The dorsal scoli 
are black with large black (approx. 0.7 mm) tumescent bases. The supralateral, lateral, 
and anal scoli are also black but lack the tumid bases. Dorsal and supralateral scoli 
vary between 3-5 mm, the anal and lateral scoli are 3 mm. Sublateral scoli Al—A8 are 
colorless and measure 1.5-2.5 mm. The prothoracic plate is large, black and elliptical, 
guages into two by a pale line through the short diameter which is at the dorsal 
midline. 


E. The Pupa 
(Fig. 2) 


The pupa is similar to that of E. lineata and other Eueides in its 
ventrally bent position and general outline. It differs in the following 
characteristics. 


Paired dorsal projections: Tl = 0.3 mm long, T2-Al = 0.2 mm, A2=0.5 mm, 
A3-A4 = 2.0 mm, A5—-A6 = 1.5 mm, A7 = 1.0 mm. Supralateral projections A3—A4 mi- 
nute, 0.1 mm. There is no evidence of bifidy and these tubercles project out at right 
angles to the body surface. All three submarginal tubercles of the wing connect to the 
wing margin by a black pigmented region. 

The general color is creamy-white marked with black spots. The dorsal scoli on 
T2-A6 are black-tipped, the rest are white. Silk pad color is white. Total length is 
16-17 mm (n = 6). It differs from other Eueides in having simple dorsal projections on 
A3 and A4 that are not very different in length from those of A5 and A6. 


DISCUSSION 


Comparison of the young stages of E. vibilia and E. lineata with 
other published accounts reveals little in the way of taxonomic sig- 


VOLUME 36, NUMBER 2 143 


nificance, but this is partly caused by the dearth of larval descriptions 
available. The pupal morphology of E. vibilia and E. lineata differs 
from other described Eueides pupae in having almost homogeneous 
dorsal projections, rather than having giant dorsal projections on A3 
and A4 as do E. isabella, E. aliphera, and E. tales Cramer (Beebe et 
al., 1960; Brown & Holzinger, 1973). Otherwise, the pupa of E. lineata 
is similar to that of E. isabella in having the dorsal projections par- 
tially bifid and by its mottled color pattern, and E. vibilia is more 
similar to that of E. aliphera in having simple spines and a black- 
spotted color pattern. 

Benson (1978) presented evidence that heliconiines partition larval 
resources. The new hostplant data presented here are concordant with 
this hypothesis. We saw no other heliconiine using Passiflora sp. nov. 
in the Sierra de Tuxtla region. New growth and old leaves on P. 
pittieri are very different resources with respect to toughness, avail- 
ability through time, and type and abundance of predators (Longino, 
unpub. data). Two allopatric species of Heliconius in Costa Rica feed 
on the new growth: H. hewitsoni on the Pacific side, and H. sapho 
Drury on the Atlantic side. E. vibilia feeds on the old leaves. _ 

The possibility of larval mimicry is intriguing. At San Pedrillo three 
heliconiine species have gregarious larvae that look extraordinarily 
similar from a human perspective: E. vibilia, H. hewitsoni, and H. 
sara (Fab.). Their yellow and black larval color pattern is very differ- 
ent from all the solitarily feeding heliconiine larvae. Why the gregar- 
ious larvae should share one pattern and the solitary larvae have other 
patterns is not easily explained. One possibility is that gregarious and 
solitary larvae have exclusive sets of predators; thus, there is no se- 
lection for a common pattern. We have some data suggesting that 
vespids and predacious pentatomids are more detrimental to gregar- 
ious than to solitary larvae, but there is no indication that these pred- 
ators are the least bit deterred by toxins or other defenses of the 
larvae. Alternatively, what is obvious to the human eye may be cryptic 
to a vespid or pentatomid eye, and the larvae may be converging on 
the pattern most cryptic to these generalized predators. E. vibilia and 
H. hewitsoni larvae occur at very low densities, but by occurring on 
the same hostplant they are, in effect, being concentrated with respect 
to the predators they experience. Thus, there could be selection for 
convergence to a common pattern if, when a bird ate one yellow 
caterpillar on a plant, it avoided all other yellow caterpillars on that 
plant. H. sara, however, is often much more abundant in a habitat, 
feeding on a more common, weedy Passiflora. Possibly, H. sara orig- 
inally evolved the yellow and black pattern as a general aposematic 
display, and they were abundant enough to provide a community- 


144 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


wide selective force for the evolution of similar patterns in the rare, 
gregariously feeding heliconiines. 


ACKNOWLEDGMENTS 


We thank Alma Solis for finding the first E. lineata at the Mexico site. We are very 
grateful to the personnel at Corcovado for allowing us to work in their park. Phil 
DeVries and L. E. Gilbert provided useful discussion and comments on the manuscript. 
The work in Mexico was partially funded by M. C. Singer and L. E. Gilbert through 
the Dept. of Zoology, Univ. of Texas, Austin. This work was ancillary to research on 
Heliconius biology funded by NSF grant, DEB-7906033, directed by L. E. Gilbert. 


LITERATURE CITED 


BEEBE, W., J. CRANE & H. FLEMING. 1960. A comparison of eggs, larvae and pupae 
in fourteen species of heliconiine butterflies from Trinidad, W. I. Zoologica, N.Y. 
45:111-153. 

BENSON, W. W. 1978. Resource partitioning in passion vine butterflies. Evolution 
32:493-518. 

BENSON, W. W., K. S. BROWN & L. E. GILBERT. 1976. Coevolution of plants and 
herbivores: passion flower butterflies. Evolution 29:659-680. 

BROWER, L. P., J. V. Z. BROWER & C. T. COLLINS. 1963. Experimental studies of 
mimicry. 7. Relative palatability and Mullerian mimicry among Neotropical but- 
terflies of the subfamily Heliconiinae. Zoologica, N.Y. 48:65—84. 

Brown, K. S. 1976. Geographical patterns of evolution in Neotropical Lepidoptera. 
Systematics and derivation of known and new Heliconiini (Nymphalidae: Nym- 
phalinae). J. Entomol. (B) 44:201-242. 

1979. Ecologia geografica e evolucao nas florestas Neotropicais. Ph.D. disser- 

tation, Universidade Estadual de Campinas, Brazil. 

1981. The biology of Heliconius and related genera. Ann. Rev. Entomol. 
26:427-456. 

Brown, K. S. & H. HOLZINGER. 1973. The Heliconians of Brazil (Lepidoptera: Nym- 
phalidae). Part IV. Systematics and biology of Eueides tales Cramer, with descrip- 
tion of a new subspecies from Venezuela. Zeit. Arbeitsgemeinsch. Osterr. Entomol. 
24:44-65. 

Brown, K. S. & O. H. H. MIELKE. 1972. The Holiconians of Brazil (Lepidoptera: 
Nymphalidae). Part IJ. Introduction and general comments, with a supplementary 
revision of the tribe. Zoologica, N.Y. 57:1—40. 

EMSLEY, M. G. 1963. A morphological study of imagine Heliconiinae (Lep., Nym- 
phalidae) with a consideration of the evolutionary relationships within the group. 
Zoologica, N.Y. 48:85-130. 

1964. The geographical distribution of the color-pattern components of Heli- 

conius erato and Heliconius melpomene with genetical evidence for the systematic 

relationships between the two species. Zoologica, N.Y. 49:245-286. 

1965. Speciation in Heliconius (Lep., Nymphalidae): morphology and geo- 
graphic distribution. Zoologica, N.Y. 50:191—254. 

GILBERT, L. E. 1975. Ecological consequences of coevolved mutualism between 
butterflies and plants. In Coevolution of Animals and Plants. L. E. Gilbert and P. 
R. Raven, eds. Univ. of Texas, Austin, p. 210-240. 

Lewis, H. L. 1974. Butterflies of the World. Harrap, London. 

Ross, G. N. 1967. A distributional study of the butterflies of the Sierra de Tuxtla in 
Veracruz, Mexico. Ph.D. dissertation, Dept. Entomol., Louisiana State Agr. and 
Mech. College. 

SMART, P. 1976. The Illustrated Encyclopedia of the Butterfly World. Hamlyn, Lon- 
don. 

SMILEY, J. T. 1978. Plant chemistry and the evolution of host specificity: new evi- 
dence from Heliconius and Passiflora. Science 201:745—-747. 


Journal of the Lepidopterists’ Society 
36(2), 1982, 145-147 


FOSSIL LEAF-MINES OF BUCCULATRIX (LYONETIIDAE) ON 
ZELKOVA (ULMACEAE) FROM FLORISSANT, COLORADO 


PAUL A. OPLER 


Office of Endangered Species, U.S. Fish and Wildlife Service, 
Department of the Interior, Washington, D.C. 20240 


ABSTRACT. A fossil leaf of Zelkova from the Florrisant formation was found to 
contain the mine of a species of Bucculatrix. This is compared to mines of extant 
species of Bucculatrix of the ulmifoliae-group. 


Leaf-mining insects, whose larvae feed between the cell layers of 
individual leaves, are found in four orders, i.e., Hymenoptera, Dip- 
tera, Coleoptera, and Lepidoptera (Needham et al., 1928). Among the 
Lepidoptera, the habit is most widespread among primitive superfam- 
ilies. Some families are composed entirely of leaf-miners, e.g., Erio- 
craniidae, Gracillariidae, Lyonetiidae, while others have both leaf- 
mining and external feeding representatives, e.g., Incurvariidae, 
Gelechiidae (Opler, 1974). Leaf-mining larvae feed in a stereotyped, 
conservative manner, and their workings may be readily identified to 
family, genus and even species, even after the responsible inhabitant 
has departed. 

The discovery of identifiable mines on finely preserved fossil leaves 
has allowed biologists not only to gain insight into the evolution of 
Lepidoptera, whose fossil record is scant, but to trace back specific 
insect-host plant relationships into geological time (Lewis, 1969; Opler, 
1973, 1974; Hickey & Hodges, 1975). 

Opler (1973, 1974) provided evidence from fossil mines that several 
oak leaf-mining moths from western North America have probably 
survived virtually unchanged since the late Miocene epoch (18 mil- 
lion years b.p.). Subsequently, Hickey and Hodges (1975) reported 
that a Phyllocnistis (Phyllocnistidae) fossil mine on Populus of Eocene 
age was clearly not the same species as any modern Populus-feeding 
Phyllocnistis. 

The present report is of Buccalatrix mines found on fossil leaves 
of Zelkova drymeja (Ulmaceae) (Fig. 1) from the Florissant formation 
in central Colorado, which is Oligocene age (30 million years b.p.). 
The host genus (Zelkova) has long since vanished from North America 
and today occurs only in temperate Eurasia. The mines are dissimilar 
to those made by any living North American species (Braun, 1963), 
but are recognizably similar to, but not conspecific with, Bucculatrix 
ulmifoliae Hg., which feeds on Ulmus in Europe and by Bucculatrix 
ulmicola Kuzn., which feeds on Zelkova in eastern Europe (Fig. 2). 

This discovery is another important piece of evidence which dem- 


146 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


FP ssh Bon a 
"Phinda pale * ay 
he. ott oe, A 
i > i vy 
' . + i. 
as ee “gl: 
POR GB ig OF 


ie £ v : Eg wx - a - > 
is) ee?” stair. Ses 
Fics. 1&2. Bucculatrix leaf mines. 1, fossil leaf of Zelkova drymeja from Florrisant 


formation with Bucculatrix mine (photograph by J. A. Powell); 2, drawing of B. ul- 
micola mines on Zelkova sp. from Rodini, Rhodos, Austria (drawing by G. Deshka). 


onstrates the historical fidelity of host relationships among the Buc- 
culatrix ulmifoliae species group with Ulmaceae for at least 30 x 10° 
years. Whether the descendents of the Florissant moths eventually 
shifted their distribution to Eurasia as antecedents of the Bucculatrix 
ulmifoliae group or whether the Florissant mines represented an in- 
vasion from Eurasia which has since died out is indeterminate. 


ACKNOWLEDGMENTS 


Jerry A. Powell provided encouragement and the photograph upon which Fig. | is 
based, while Gerfried Deshka provided the drawing of Bucculatrix ulmicola mines for 
Fig. 2. Both gentlemen are gratefully acknowledged. I also thank H. D. McGinitie for 
determining the Zelkova impression from Florissant. 


LITERATURE CITED 


BRAUN, A. F. 1963. The genus Bucculatrix in America north of Mexico. Mem. Am. 
Entomol. 18:1—208. 

HIckEY, L. J. & R. W. HODGES. 1975. Lepidopteran leaf mine from the early Eocene 
Wind River formation of northwestern Wyoming. Science 189:718-720. 

LEwIs, S. E. 1969. Lepidopterous larval mining on an oak (?) leaf from the Latah 
Formation (Miocene) of eastern Washington. Ann. Entomol. Soc. Amer. 62:1210. 


_ VOLUME 36, NUMBER 2 147 
| 


NEEDHAM, J. G., S. W. FRosT & B. H. TOTHILL. 1928. Leaf-mining Insects. Williams 
and Wilkins Co., Baltimore. 351 pp. 


OPLER, P. A. 1973. Fossil lepidopterous leaf mines demonstrate the age of some 
insect-plant relationships. Science 179:1321-1323. 

1974. Biology, ecology, and host specificity of Microlepidoptera associated 

with Quercus agrifolia (Fagaceae). Univ. California Publ. Entomol. 75:1-83. 


Journal of the Lepidopterists’ Society 
36(2), 1982, 148-152 


LIZARD PREDATION ON TROPICAL BUTTERFLIES 


PAUL R. EHRLICH AND ANNE H. EHRLICH 
Department of Biological Sciences, Stanford University, Stanford, California 94305 


ABSTRACT. Iguanid lizards at Iguacu Falls, Brazil appear to make butterflies a 
major component of their diets. They both stalk sitting individuals and leap into the 
air to capture ones in flight. Butterfly species seem to be attacked differentially. These 
observations support the widespread assumption that lizards can be involved as selec- 
tive agents in the evolution of butterfly color patterns and behavior. 


Butterflies have been prominent in the development of ideas about 
protective and warning coloration and mimicry (e.g., Cott, 1940; J. 
Brower, 1958; M. Rothschild, 1972), and the dynamics of natural pop- 
ulations (Ford & Ford, 1930; Ehrlich et al., 1975). In spite of the 
crucial role that predation on adults must play in evolution of defen- 
sive coloration and may play in population dynamics, there is re- 
markably little information on predation on adult butterflies in nature. 
This lack is all the more striking, considering the large numbers of 
people who collect butterflies and the abundant indirect evidence 
from bird beak and lizard jaw marks on butterfly wings (e.g., Carpen- 
ter, 1937; Shapiro, 1974) that adult butterflies are quite frequently 
attacked. 

Published field observations of predation on butterflies deal almost 
exclusively with the attacks of birds and consist largely of accounts 
of individual attacks (Fryer, 1913). Observations of natural predation 
by lizards are very rare, although “birds and lizards have long been 
considered to be the major selective agents responsible for the ex- 
treme diversity of unpalatable and mimetic forms of butterflies in 
nature’ (Boyden, 1976). The following observations confirm the po- 
tential ability of lizards to place powerful selection pressures on but- 
terfly populations. 

A group of about seven iguanid lizards, Tropidurus torquatus (Wied), 
were observed on rocks adjacent to a walkway below the brink of 
Iguacu Falls in southwestern Brazil on 26 November 1980. The larg- 
est had a snout-vent length of about 15 cm; the others were about 10 
cm or slightly smaller. While we were watching, a small, colorful 
nymphaline butterfly, Callicore hydaspes Drury, flew by about 50 cm 
above the lizards, several of which turned their heads to watch it pass. 
A few minutes later a small nymphaline (possibly Dynamine arte- 
misia Felder) landed on the rocks about 15 cm from a lizard, which 
lunged at it, captured it, and ate it. 

It subsequently proved possible to make roughly five person hours 
of undisturbed observations in sunny weather in the late mornings of 


VOLUME 36, NUMBER 2 149 


26 and 27 November. During that period we saw hundreds of lizard 
“reactions” to butterflies—flight-following with the head, short move- 
ments in the direction of a butterfly that had landed, or prolonged 
gradual stalking of sitting butterflies (Fig. 1). About 75 clear attacks 
were observed, consisting of a lunge that carried the lizard to or past 
the position previously occupied by the butterfly (Fig. 2) or a leap 
clear of the ground in the direction of a flying butterfly. Fifteen but- 
terflies were captured and devoured. The butterflies eaten were the 
Dynamine, 1 Eunica margarita Godart, 6 Callicore hydaspes, 2 Mar- 
pesia chiron Felder, 1 M. petreus Cramer, and 1 Dione juno Cramer 
(all Nymphalidae: Nymphalinae); 1 yellowish-white pierid (possibly 
a female Phoebis statira Cramer); 1 small bluish skipper (Hesperioi- 
dea), and 1 large, powerful skipper (possibly an Astraptes or Pyrrho- 
pyge). 

The response of the lizards to different butterfly species was quite 
variable. They showed the greatest interest in C. hydaspes, which 
was also the commonest in the area. Its appearance in flight invariably 
invoked a reaction, even at a distance of a meter or more. When other 
butterflies passed by, however, very often there was no movement on 
the part of the lizards. Many of the butterflies landed on a small sandy 
patch next to the rocky area occupied by the lizards and showed clas- 
sic “puddling” behavior (Fig. 1), probing the sand with their probos- 
cides and dripping water from the anus—presumably acquiring salts 
(Arms et al., 1974), in this case possibly from lizard droppings. Gen- 
erally lizards would stalk these butterflies until they were within 10-20 
cm and then lunge at them. Butterflies that landed on the rock itself 
tended to elicit more rapid attacks, tempting one to speculate that the 
lizards had learned that butterflies not puddling were less likely to 
remain in place for an extended period. It also seemed that the pres- 
ence of another nearby lizard prompted more immediate attack. 

Leaps at passing butterflies were surprisingly frequent and roughly 
as successful as surface attacks (about | in 5). The Dione and one C. 
hydaspes were captured in mid-air, as was one large skipper, which, 
however, managed to wrench itself free and escape after the lizard 
had returned to earth. Lizards in other circumstances may attempt to 
catch flying butterflies—lizard jaw marks on only one wing may be 
evidence of this (L. Gilbert, pers. comm.) since butterflies normally 
sit with their wings held together over their backs (Fig. 1). Lizards 
have also been observed to leap clear of the ground to catch dragon- 
flies on the wing (T. Schoener, pers. comm.). In experimental work 
on the palatability of butterflies to teiid lizards (Ameiva ameiva L.), 
Boyden (1976) found that when tethered butterflies “got stuck in tall 
grass above the lizard’s head ... the Ameiva would frequently jump 


150 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


, af ved aa { by é at - » . i ™ Ve 
Fics. 1-2. 1, Tropidurus torquatus stalking two Marpesia chiron (to the right of 
the lizard’s head) and one M. petreus (below the lizard’s head). Butterfly wingspreads 
approximately 50 mm. 2, an unsuccessful lunge, a moment after the photo in Fig. 1 
was taken. Note that the M. petreus in the upper right (tip of its wing barely visible 
in Fig. 1) remains undisturbed. 


VOLUME 36, NUMBER 2 151 


distances greater than 0.4 m off the ground to attack the butterfly, pull 
it to the ground, and eat it.” Nonetheless 13 of 15 Tropidurus captures 
observed by us were of sitting butterflies. This, not surprisingly, con- 
trasts with the pattern of bird attacks, where more attacks seem to be 
aerial (Collenette, 1935; Carpenter, 1937; Bowers & Wiernasz, 1979). 
The vast majority of attacks observed by Shapiro (1974), however, 
were on sitting butterflies, and recently, evidence of heavy bird pre- 
dation on resting Euphydryas chalcedona has been found (D. M. 
Bowers and I. L. Brown, in preparation). 

Every butterfly captured at Iguacu was completely devoured, so 
that no evidence of predation in the form of severed wings remained. 
In the process of swallowing the captured Dione, the lizard broke off 
a large piece of the butterfly’s hindwing. After the rest of the butterfly 
was consumed, the lizard picked up the remaining piece of wing and 
swallowed it too. In contrast, wings are often removed by birds before 
the body is eaten (Collenette, 1935; Carpenter, 1937), and in at least 
one case of observed lizard attack on a temperate zone butterfly (Va- 
nessa cardui L.), an iguanid (Sceloporus graciosus B.-G.) beat the 
butterfly against the ground to remove its wings before swallowing 
the body (Knowlton, 1953). 

The Dione was the only butterfly attacked that, on the basis of its 
taxonomic affinities, might reasonably be expected to be at least some- 
what unpalatable. Brower et al. (1963) found that close relatives of 
the Dione in the Heliconiini, Dryas julia Fabricius and Agraulis va- 
nillae L., were unpalatable to silverbeak tanagers, although less so 
than members of the genus Heliconius. Several Heliconius passed 
within 1 m of the lizards we were observing but did not elicit the 
reactions that the smaller Callicore invariably did at the same dis- 
tance. Boyden’s work and greenhouse observations (L. Gilbert, pers. 
comm.) indicate that lizards find certain butterflies unpalatable and 
can learn to avoid them, and this seems a reasonable explanation for 
the behavior of Tropidurus toward Heliconius. 

Although the lizards were also observed snapping at and catching 
small flies, during our observations butterflies were occupying most 
of their attention and in volume made up the vast majority of their 
intake. Butterflies are very abundant at Iguacu because of extremely 
extensive forest-edge situations created by the falls and the facilities 
of Iguacu National Park. They seemed especially common along the 
observation trails, frequently landing on wooden and metal handrails, 
presumably attracted by the salts left by sweating tourists. Lizards 
were abundant in precisely the same areas, and other insects were 
not conspicuous. 

These observations indicate that, at least for some species such as 


152 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Callicore hydaspes, lizards may be significantly able to affect popu- 
lation size through predation on the adults. Since they evidently dif- 
ferentiate between butterfly species under natural conditions, they 
may well influence the evolution of butterfly color patterns and be- 
havior. 


ACKNOWLEDGMENTS 


We thank M. D. Bowers, R. W. Holm, H. A. Mooney, D. D. Murphy and M. C. Singer 
for criticism of the manuscript. Keith Brown was helpful in identifying the butterflies; 
Richard Etheridge kindly identified the lizards. We are grateful to José and Sonia 
Rotenberg for support in the field. 


LITERATURE CITED 


ARMS, K., P. FEENY & R. C. LEDERHOUSE. 1974. Sodium: Stimulus for puddling 
behavior by tiger swallowtail butterflies, Papilio glaucus. Science 185:372-374. 

BowERs, M. D. & D. C. WIERNASZ. 1979. Avian predation on the palatable butterfly 
Cercyonis pegala (Satyridae). Ecol. Entomol. 4:205-209. 

BOYDEN, T. C. 1976. Butterfly palatability and mimicry: Experiments with Ameiva 
lizards. Evol. 30:73-81. 

BROWER, J. V. Z. 1958. Experimental studies of mimicry in some North American 
butterflies. Part I. The Monarch, Danaus plexippus and Viceroy, Limenitis ar- 
chippus archippus. Evol. 12:32-47. 

BROWER, L. P., J. V. Z. BROWER & C. T. COLLINS. 1963. Experimental studies of 
mimicry. 7. Relative palatability and Mullerian mimicry among neotropical but- 
terflies of the subfamily Heliconiinae. Zoologica 48:65-84. 

CARPENTER, G. D.H. 1937. Further evidence that birds do attack and eat butterflies. 
Proc. Zool. Soc. Lond. (A). 107:223-247. 

COLLENETTE, C. L. 1935. Notes concerning attacks by British birds on butterflies. 
Proc. Zool. Soc. Lond. 1935:200-217. 

Cott, H. B. 1940. Adaptive coloration in animals. Methuen, Lontiont 508 pp. 
EHRLICH, P. R., R. R. WHITE, M. C. SINGER, S. W. MCKECHNIE & L. E. GILBERT. 
1975. Checkerspot butterflies: A historical perspective. Science 188:221—228. 
ForpD, H. D. & E. B. Forp. 1930. Fluctuation in numbers and its influence on vari- 

ation in Melitaea aurinia. Trans. R. Entomol. Soc. Lond. 78:345-351. 

FRYER, J. C. F. 1913. Field-observations on the enemies of butterflies in Ceylon. 
Proc. Zool. Soc. Lond. 2:613-619. 

KNOWLTON, G. F. 1953. Predators of Vanessa cardui. Lepid. News 7:55. 

ROTHSCHILD, M. 1972. Some observations on the relationship between plants, toxic 
insects and birds. Pp. 1-12 in J. B. Harborne (ed.), Phytochemical Ecology, Aca- 
demic Press, London. 272 pp. 

SHAPIRO, A. M. 1974. Beak-mark frequency as an index of seasonal predation inten- 
sity on common butterflies. Amer. Nat. 108:229-232. 


Journal of the Lepidopterists’ Society 
36(2), 1982, 153 


GENERAL NOTES 


DISTINCTIVENESS OF MEGISTO C. CYMELA AND 
M. C. VIOLA (SATYRIDAE) 


Megisto cymela Cramer is remarkable for its lack of geographic variation over a range 
that extends from Manitoba and Quebec to the Gulf Coast. The subspecies M. c. viola 
Maynard, described from Florida, is highly distinctive, differing chiefly by its larger 
size and rich coloration on the ventral hindwing. Most references (e.g., A. B. Klots, 
1951, A Field Guide to the Butterflies, Houghton Mifflin, Boston, 349 pp.; W. H. Howe, 
1975, Butterflies of North America, Doubleday, New York, 633 pp.) consider that the 
name viola refers to all peninsular Florida populations and that M. c. cymela and M. 
c. viola blend phenotypically in southern Georgia and northern Florida. 

An examination by me of series in The Florida State Collection of Arthropods, 
Gainesville, Florida, indicates that this is not the case. There is no sign of a phenotypic 
“blend zone,” and populations of both typical M. c. cymela and M. c. viola exist in 
northern and central Florida. In addition, on the basis of data labels in The Florida 
State Collection, M. c. viola exists apparently sympatrically with M. c. cymela in south- 
ern Louisiana (Weyanoke, West Feliciana Parish) and Arkansas (Little Rock, Pulaski 
Co.). Moreover, in Florida M. c. viola appears strictly univoltine with a flight from late 
March to May. In the same general area M. c. cymela may have up to four broods with 
flights in April (Shalimar, Okaloosa Co.; Alachua Co.), July (Alachua Co.), October 
(Gainesvilla, Alachua Co.), and December (Sebring, Highlands Co.). The two entities 
thus may well be separate species. 

Larvae of Pennsylvania M. c. cymela and Florida M. c. viola differ in life history in 
the laboratory. Although the pattern of larval markings is similar, larvae of M. c. cymela 
are a much darker shade of brown than M. c. viola. Broods of M. c. cymela derived 
from univoltine populations in Allegheny and Fayette Cos., Pennsylvania, develop 
without diapause and emerge as adults in 90 to 100 days when reared under conditions 
of 27°C days, 24°C nights and 16 hr light/24 hr. This correlates well with the three to 
four months between flights of M. c. cymela in Florida. Under the same conditions, 
however, growth of M.c. viola larvae from a Gainesville population differs greatly. 
Larvae hatched from eggs in early April 1979 grew at a very slow but steady rate and 
pupated and eclosed as adults in late February and early March 1980. This also cor- 
relates well with the flight time of M. c. viola in Florida if the cooler winter temper- 
atures in nature are taken into consideration. 

Larvae of both entities were reared on potted Poa pratensis L., a grass native to the 
northern U.S. This could be a natural foodplant of M. c. cymela, but M. c. viola occurs 
south of its natural range. Nevertheless, M. c. viola larvae fed freely and produced 
adults of a size comparable to wild material. I consider it highly unlikely that differ- 
ences in development time were related to foodplant suitability. 

Collectors in the Gulf Coast states should watch for localities where M. c. cymela 
and M. c. viola are sympatric and attempt to gather data on possible ecological differ- 
ences. 


CHARLES G. OLIVER, R.D. 1, Box 78, Scottdale, Pennsylvania 15683. 


Journal of the Lepidopterists’ Society 
36(2), 1982, 154-155 


LONG MATING FLIGHTS BY MALE 
HYALOPHORA CECROPIA (L.) (SATURNIDAE) 


Distances from which male moths can find the pheromone-emitting females have 
been determined for a few species by releasing marked males and recapturing them 
in traps baited with females. For example, Rau and Rau (1929, Trans. Acad. Sci. St. 
Louis 26:81-221) recovered about 11% of the male cecropia moths released 4.8 km 
from their trap. Distances of over 10 km have been recorded with other species (Shorey, 
1976, Animal Communication by Pheromones, Academic Press, New York). (Also see 
below.) 

These and similarly obtained data (including ours) are not a measure of the distance 
over which males can perceive the sex attractant pheromone. The males’ flights will, 
of course, include random searching as well as directed movement to the phero- 
mone-probably via anemotaxis (Kennedy, 1977, pp. 67-91 In: Chemical Control of 
Insect Behavior, H. H. Shorey and J. J. McKelvey, Jr., eds., John Wiley, New York). 
Neither do such data represent the entire distance flown by the males. Their actual 
flight paths are unknown, but are probably almost always longer than the straight line 
distance between the release and recapture points. Furthermore, data such as these 
are only minimum estimates of the straight line distance over which males can find 
females. If males are released at even greater distances, some may eventually be re- 
covered; Toliver and Jeffords (1981, J. Lepid. Soc., in press) recaptured a male Cal- 
losamia promethea (Drury) 36.5 km from the release point. 

Exceptionally long mating flights may usually be uncommon events, but they are 
probably important in promoting gene flow to distant populations, and they may make 
it possible for populations to survive at very low densities. Since information on long 
mating flights is scarce, we here present data that were obtained in the course of another 
study. 

We used three traps baited with virgin female cecropia (Sternburg and Waldbauer, 
1969, Ann. Entomol. Soc. Amer. 62: 1422-1429) to catch wild males in, and near, the 
contiguous cities of Champaign and Urbana, Illinois, from 14 May to 10 July. One trap 
was at the home of JGS near the eastern edge of the cities, another was at the home 
of GPW near the western edge of the cities, and the third was east of the cities at 
Trelease Woods. Straight line distances between the traps appear in Table 1. Males 
came to the traps just before dawn. Later they were consecutively numbered on the 
underside of the hindwing with a felt pen—a different color for each trap—and released 
at the capture site. They dispersed, but we don’t know how far they went. However, 
they usually flew out of sight even in daylight, and presumably made another dispersal 
flight at dusk, several hours before the females again emitted pheromone (Waldbauer 
and Sternburg, 1979, Amer. Midland Nat. 102:204—208). 

We caught 1069 wild males and recaptured 390 (36.5%) of them one or more times. 
Twenty (5.1%) of the latter were recaptured at a more distant trap (Table 1), including 
one reared male (not in Table 1) released at a park in Champaign and recaptured 10.1 
km away at Trelease. 

Wind direction and velocity are important factors in the dispersal of pheromones. 
They determine the direction in which the male will fly and affect the distance that 
he can fly. We cannot determine the wind conditions that prevailed while the males 
made their way to the recapture sites. Wind direction and velocity may vary from hour 
to hour, and all but two of the nineteen recaptured males in question had from two to 
eight days to reach the recapture sites (Table 1). Thus, the winds almost certainly varied 
as these males moved toward the traps on any one of several nights or on some com- 
bination of two or more nights. Weather records for our area include the mean daily 
wind velocity and direction; these generally varied from day to day between the day 
of release and the day of recapture. 

The frequency of occurrence of long flights may change with the noneleeron density 
of the wild moths. At lower densities than prevailed during our experiment, more long 
flights should occur because wandering males will be less likely to encounter the aerial 


VOLUME 36, NUMBER 2 155 


TABLE 1. Number of male Hyalophora cecropia recaptured at sites other than the 
release point, the distance between these points, and the number of days that elapsed 
between release and recapture. 


Trap at which male was: 
Distance (km) Days (or range) 


Released Recaptured between traps No. males between captures 
Gs GPW 6.8 8 3-8 
GPW JGS 6.8 8 1-7 

Trelease JGS 6.8 1 1 
JGS Trelease 6.8 1 2 
GPW Trelease 1A) 1 8 


pheromone trail of a wild female before encoutering a trail from one of the traps. 
Similarly, in natural situations (without traps), the average distance flown by males 
searching for females should increase as the population density decreases. Our data 
(Table 1) suggest that for male cecropia moths these distances may be very long, and 
that reproduction could, therefore, continue even at very low population densities. 


G. P. WALDBAUER & J. G. STERNBURG, Department of Entomology, 320 Morrill 
Hall, University of Illinois, Urbana, Illinois 61801. 


Journal of the Lepidopterists’ Society 
36(2), 1982, 155-157 


HINDTIBIAL DEFENSIVE SPURS IN THE NEOTROPICAL 
SPHINX MOTH AMPLYPTERUS GANNASCUS? 


A noticeable morphological feature of most adult Sphingidae is the double pair of 
elongate spurs on the tibia of the hindlegs (W. Rothschild and K. Jordan, 1903, A 
Revision of the Lepidopterous Family Sphingidae, Novitates Zool., Vol. 9, Suppl.). 
There is usually a proximal and terminal pair of these spurs, which are modified spines. 
The functional role, if any, of these structures is unknown. However, they are con- 
spicuously long and rigid in some sphingid taxa. In this note I wish to suggest a possible 
defensive role of these hindtibial spurs in sphingids, based upon my being jabbed by 
the spurs of Amplypterus gannascus (Stoll) to the point of considerable bleeding. 

On the evening of 19 June 1980 at about 1600 hours, I collected several freshly- 
eclosed adults of various sphingids resting on the lighted wall of a “cacao beneficio” 
building at “Finca Experimental La Lola,” a large cacao plantation along the railroad 
line connecting Guapiles with the Caribbean port city of Limon, Limon Province, Costa 
Rica. Upon picking up one of the largest specimens, I suddenly felt a very painful stab 
into one of my fingers. At first I thought it was a wasp sting, thinking that I had 
inadvertently picked up a wasp in the shadows along with the moth. A copious flow 
of blood told me that I had been jabbed with something very sharp. The moth turned 
out to be A. gannascus and close examination revealed very long (10 mm) and stout 
hindtibial spurs (Fig. 1) capable of piercing soft tissues without hindrance. The spurs 
were present only on the hindtibiae of the moth. 

Amplypterus gannascus is widespread throughout Central and South America and 
the Caribbean (A. Seitz, 1924, Macrolepidoptera of the World, Vol. 5, American Rho- 
palocera, Stuttgart) and it is one of the larger sphingids attracted to lights in lowland 


156 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Fic. 1. Above: Amplypterus gannascus from Finca Experimental La Lola, in south- 
eastern Costa Rica, dorsal view; Below: hindtibial spurs of A. gannascus pointing 
upward and just to the left of the tarsal region. 


tropical rain forest areas. Being a large insect, the moth, as well as other sphingids, 
may be a target for predatory vertebrates that forage at lights in the tropics. The grass 
and lower vegetation at the base of lighted walls is often littered with large toads that 
readily could take moths in the act of alighting. Certain species of bats may also take 
adult moths as they fly around lights. Both groups of insectivorous predators are abun- 


VOLUME 36, NUMBER 2 157 


dant in the tropics. Although sphingid moths commonly accumulate at lights along with 
many other insects in the wet tropics, the kind of defense suggested here would be 
adaptive when away from lights as well. Moths flying through darkness might be picked 
off by foraging bats. Some Neotropical insectivorous bats readily respond to wing- 
flapping noises from large moths confined to the same cages (M. D. Tuttle, pers. comm.). 
Successful capture of a large sphingid results in the predator obtaining a large protein- 
and lipid-rich food morsel. 

The presence of hindtibial spurs of varying size in sphingids may represent an ad- 
aptation to defense against predators. When jabbed with the spurs, my reaction was to 
immediately release the captured insect, permitting its escape. Similar behavioral re- 
sponses might occur if spurs can successfully lodge in soft tissues around or just inside 
the mouth of a predator. Alternatively, these pronounced spurs may have little or no 
direct defensive function per se, and perhaps are non-functional, or are used in other 
activities such as courtship or feeding, but, secondarily can be used opportunistically 
in defense. Closer scrutiny of the functional role of these spurs, in sphingids in general, 
is needed. Studies, including analyses of distribution among sexes within a species, 
spur sizes, and frequency of occurrence in tropical and extra-tropical taxa, should be 
done. 

I thank Susan S. Borkin for technical assistance and Dr. Merlin D. Tuttle for sharing 
with me his knowledge of moth predation by Neotropical bats. 


ALLEN M. YOUNG, Section of Invertebrate Zoology, Milwaukee Public Museum, 
Milwaukee, Wisconsin 53233. 


Journal of the Lepidopterists’ Society 
36(2), 1982, 157-158 


TWO SPECIES OF SKIPPERS COLLECTED AT ANTIFREEZE-FILLED 
PITFALL TRAPS IN ARIZONA 


During a research trip to the Southwestern Research Station, American Museum of 
Natural History, Portal, Arizona 85632 in 1978, the senior author was able to collect 
hundreds of skippers in antifreeze-filled pitfall traps. The skippers were subsequently 
determined by the junior author as Atrytonopsis python (Edwards) and A. deva (Ed- 
wards). 

Four pitfall traps (plastic cottage cheese containers) were placed, flush with the 
ground, 10’—20’ apart around a pond that had a heavy red algal bloom. The traps were 
placed 3’—4' from the water’s edge with the intention of collecting ground dwelling 
beetles near the water’s edge. The traps were % filled with Dowguard® antifreeze for 
specimen preservation. The traps were checked every 2-3 days, emptied and new 
antifreeze put in to bring the trap level back up to % full. 

During the month of June, hundreds of the two above mentioned skippers were 
found in the traps, preserved in the antifreeze. A select number of the skippers were 
removed from the antifreeze traps and taken to the lab. They were carefully washed 
with 75% ETOH and placed on paper towel to dry. When the alcohol absorbed by the 
towel had dried, the specimens were pinned and spread. No adverse effects on scale 
coloration of the specimens were noted. 

To the author's knowledge, this is the only known record of skippers being attracted 
to antifreeze. As ethylene glycol is present in both the antifreeze and the insects (Somme, 
1964. Canad. Jour. Zool. 42:87-101), it may be acting as a “cue” to attract the 
skippers to the traps. Skippers of this genus generally are attracted to flowers, some- 
times in swarms, according to Howe (1975, The Butterflies of North America, Double- 
day & Co., Garden City, N.Y.). Use of this type of artificial lure may be a useful method 


158 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


of collecting these and other Atrytonopsis species. Specimens of both species have 
been deposited in the Michigan State University collection. 


DUANE J. FLYNN, Department of Entomology, Michigan State University, East Lan- 
sing, Michigan 48824 & MOGENS C. NIELSEN, 3415 Overlea Dr., Lansing, Michigan 
48917. 


Journal of the Lepidopterists’ Society 
36(2), 1982, 158-159 


A ONE-FOURTH GYNANDROMORPH OF AGRIADES RUSTICA RUSTICA 
(EDWARDS) FROM WYOMING (LYCAENIDAE) 


On a recent collecting trip to the Bighorn Mountains of Wyoming, an unusual single 
specimen of A. r. rustica (W. H. Edwards) was collected. The coloration of the wings, 
together with their relative size, clearly shows the left hindwing to be male, with the 
other three wings being female. In this respect it is similar to the specimen of Strymon 
bazochii (Godart) illustrated by Riotte [1978 (1979), J. Res. Lepid. 17(1):17-18.]. Ex- 
amination of the external features suggests that the abdomen is that of a female spec- 
imen. While no other gynandromorphic Plebejinae are known to me from the Nearctic 
region, Ford (1945, Butterflies, London, pp. 193-195) illustrates a number of interesting 
forms from the Palearctic region. 

Nielson (1977, J. Res. Lepid. 16(4):209-211) has pointed out that there has been a 
recent increase in publications dealing with gynandromorphism in the Lepidoptera. 


Fic. 1. Gynandromorph of Agriades rustica rustica from Wyoming. (Photo credit: 
Steve Lewis) 


VOLUME 36, NUMBER 2 159 


Examination of his literature citations, as well as those of Perkins and Perkins (1972, 
J. Res. Lepid. 11(3):195-196) indicates that the last twenty years have seen the ap- 
pearance of a larger proportion of papers on that subject than in the decades preceding 
1960. Whether or not this indicates an increase in the frequency of gynandromorphism 
or merely an increased awareness in the occurrence of the phenomenon is still a matter 
of speculation. It seems, however, that the interests of earlier collectors and authors 
for varieties and aberrations would have disclosed more examples from the Nearctic 
than are apparent at present. 

The data for the specimen illustrated are as follows: Johnson County, Wyoming, 
Bighorn Mountains, Powder River Pass, 9660’, 23 July 1980. At present, the specimen 
is in the author's collection. 


RUSSELL RAHN, 3205 W. Rochelle Road, Irving, Texas 75062. 


Journal of the Lepidopterists’ Society 
36(2), 1982, 159 


CHANGE IN STATUS OF CATOCALA ANDROMACHE RACE 
“BENJAMINI” (NOCTUIDAE) 


Few specimens of the andromache complex of Catocala were available in collections 
in the early nineteen thirties when Dr. Foster H. Benjamin and I stood together in the 
U.S. National Museum and he suggested that I describe a unique portion (benjamini) 
of the complex as a race of andromache Hy. Edw, which I subsequently did (Brower, 
S. E., 1937, Bull. Brooklyn Entomol. Soc. 32(5):185-186). The unique type of andro- 
mache is considerably worn, and at the time we were considering this problem, quite 
a few collectors seemed to feel that there were already too many specific names in the 
Catocala. C. benjamini, NEW STATUS, is more uniformly brown-gray, with broader 
more prominent lines, than others of the complex, and most specimens lack to some 
degree the greenish shade characteristic of many andromache. No polymorphism in 
benjamini is evident to me in the material at hand. When I wrote the original descrip- 
tion of the latter, the known ranges of the two entities were allopatric; however, I 
currently have data showing sympatry over portions of their ranges. 

Based on an examination of the features mentioned above relative to significant series 
of all recognized species in the andromache complex, I conclude that benjamini Brow- 
er should be elevated to species level. I am making the change at this time so the new 
status will be available for use in detailed biological studies on this complex by J. W. 
Johnson and E. Walter now awaiting publication. 


A. E. BROWER, 8 Hospital Street, Augusta, Maine 04330. 


Journal of the Lepidopterists’ Society 
36(2), 1982, 159 


A RECORD OF ITAME ABRUPTATA (GEOMETRIDAE) FROM WISCONSIN 


Hoebeke (1980, J. Lepid. Soc. 34:132) reported the first occurrence of Itame abrup- 
tata (Walker) in New York and noted that it was “not well represented in North Amer- 
ican collections,’ as cited in McGuffin’s work (1977, J. Lepid. Soc. 31:269-274). McGuffin 
included Wisconsin in the range of the foodplant, but indicated no records of the moth 
from that state. Among material I received from Irwin Leeuw and determined for me 
by F. H. Rindge, American Museum of Natural History, New York, were the following: 
one male and one female, taken 9 July 1979 in Grant Co., Wisconsin. 


BRYANT MATHER, 213 Mt. Salus Drive, Clinton, Mississippi 39056. 


Journal of the Lepidopterists’ Society 
36(2), 1982, 160 


NEW HOSTPLANT RECORDS FOR AGONOPTERIX CLEMENSELLA 
(OECOPHORIDAE) 


As is the case for most of the Oecophoridae, details of the biology of early stages of 
Agonopterix clemensella (Chambers) are poorly known. Hodges (1974, Gelechioidea 
Oecophoridae, Moths of America North of Mexico, Fascicle 6.2, E. W. Classey, Ltd., 
London) reports that “the larva has been reared from parsnip, Pastinaca sativa L. and 
undoubtedly feeds on native umbels.” Exhaustive sampling of Umbelliferae in Tomp- 
kins County, New York, during the spring and summer months of 1977 through 1979, 
revealed that A. clemensella utilizes a broad range of both native and introduced species 
(Table 1). The host list includes representatives of two subfamilies and five tribes in 
the family Umbelliferae; ten of the sixteen species are native to North America. The 
host plants occur in a variety of habitats, ranging from rich woods to waste places; 
clearly, A. clemensella is a family and not a habitat specialist. 

The only umbellifer examined that is consistently avoided by A. clemensella is 
Conium maculatum (poison hemlock); in fact, caterpillars confined to the foliage in- 
variably died. C. maculatum, however, is the sole host for A. alstroemeriana, a recently 
introduced European species (Berenbaum and Passoa, in preparation). 

Larvae can be collected throughout June; adults emerge in late June and early July, 
approximately 10-14 days after pupation. Caterpillars on each plant species were reared 
through to the adult stage to verify their identity. Identifications were made by J. 
Franclemont and R. Brown of the Department of Entomology at Cornell University; 
representative specimens are on deposit in the Cornell University Collection, Lot 1023, 
Sublot 41B. This work was supported by National Science Foundation research grant 
DEB 76-20114 to P. Feeny. 


M. BERENBAUM, Department of Entomology, Cornell University, Ithaca, New York 
14853. (Present address: Department of Entomology, 320 Morrill Hall, University of 
Illinois at Urbana-Champaign, Urbana, Illinois 61801.) 


TABLE 1. Hostplants of Agonopterix clemensella in Tompkins County, New York. 
(Species arranged according to Drude, 1898, Umbelliferae, in Die natiirlichen Pflan- 
zenfamilien 3:63-250.) | 


Saniculoideae 
Saniculeae *Sanicula gregaria Damp woods 
Apioideae 
Scandicinae *Osmorhiza longistylis Damp woods 
Carinae Apium graveolens Greenhouse 
*Zizia aptera Dry woods 
*Zizia aurea Damp woods 
*Cicuta maculata Wet meadows 
*Cryptotaenia canadensis Damp woods 
*Taenidia integerrima Dry woods 
Aegopodium podagraria Waste places 
*Sium suave Wet meadows 
Peucedaneae *Angelica atropurpurea Wet meadows 
Levisticum officinale Waste places 
Pastinaca sativa Waste places 
*Heracleum lanatum Waste places 
Heracleum mantegazzianum Waste places 
Dauceae Daucus carota Waste places 


* Species considered native to North America (according to Fernald, 1950, Gray's Manual of Botany, 8th edition, 


American Book Co., 


Date of Issue (Vol. 36, No. 2): 24 September 1982 


EDITORIAL STAFF OF THE JOURNAL 
THOMAS D. EICHLIN, Editor 


% Insect Taxonomy Laboratory 
1220 N Street 
Sacramento, California 95814 U.S.A. 


MAGDA R. PAppP, Editorial Assistant 


DOUGLAS C. FERGUSON, Associate Editor THEODORE D. SARGENT, Associate Editor 


NOTICE TO CONTRIBUTORS 


Contributions to the Journal may deal with any aspect of the collection and study of 


_ Lepidoptera. Contributors should prepare manuscripts according to the following in- 
structions. 


Abstract: A brief abstract should precede the text of all articles. 

Text: Manuscripts should be submitted in triplicate, and must be typewritten, 
entirely double-spaced, employing wide margins, on one side only of white, 8% x 11 
inch paper. Titles should be explicit and descriptive of the article's content, including 


_the family name of the subject, but must be kept as short as possible. The first mention 
_ of a plant or animal in the text should include the full scientific name, with authors 
of zoological names. Insect measurements should be given in metric units; times 
_ should be given in terms of the 24-hour clock (e.g. 0930, not 9:30 AM). Underline only 


where italics are intended. References to footnotes should be numbered consecutively, 


and the footnotes typed on a separate sheet. 


Literature Cited: References in the text of articles should be given as, Sheppard 
(1959) or (Sheppard 1959, 196la, 1961b) and all must be listed alphabetically under 
the heading LITERATURE CITED, in the following format: 


SHEPPARD, P. M. 1959. Natural selection and heredity. 2nd. ed. Hutchinson, London. 
209 pp. 

1961a. Some contributions to population genetics resulting from the study of 

the Lepidoptera. Adv. Genet. 10: 165-216. 


In the case of general notes, references should be given in the text as, Sheppard (1961, 


_ Adv. Genet. 10: 165-216) or (Sheppard 1961, Sym. R. Entomol. Soc. London 1: 23-30). 


Illustrations: All photographs and drawings should be mounted on stiff, white 
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inches are not acceptable and should be reduced photographically to that size or small- 


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Proofs: The edited manuscript and galley proofs will be mailed to the author for 


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Correspondence: Address all matters relating to the Journal to the editor. Short 
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the editor of the News: W. D. Winter, Jr., 257 Common Street, Dedham, Massachusetts 


02026 U.S.A. 


PRINTED BY THE ALLEN PRESS, INC., LAWRENCE, KANSAS 66044 U.S.A. 


CONTENTS 


FOODPLANT AND OVIPOSITION RECORDS FOR PANAMANIAN 
LYCAENIDAE AND RIODINIDAE. Robert K. Robbins & An- 
nette Aiello (han A 0 

DIFFERENTIAL GROWTH AND UTILIZATION OF THREE FOOD- 
PLANTS BY FIRST INSTAR LARVAE OF CITHERONIA REGALIS 
(SATURNIIDAE). C. Brooke Worth, Austin P. Platt J 
Thomas Fe Williams 2. a ee A 


DYSSTROMA HERSILIATA FORM “MIRANDATA’ —A RECESSIVE COL- 


OR FORM (GEOMETRIDAE). K. B. Bolte... aaa 


DATES OF SELECTED LEPIDOPTERA LITERATURE FOR THE 
WESTERN HEMISPHERE FAUNA. John B. Heppner _______ 


THE CHROMOSOMES OF A WILD SILKMOTH, ARCHAEOATTACUS 
EDWARDSII, WITH A RECORD HIGH CHROMOSOME NUMBER 
FOR SATURNIIDAE. R.C. Narang & M. L. Gupta __..-.____ 


ADDENDUM AND CORRIGENDA TO “CLASSIFICATION OF THE 
SUPERFAMILY SESIOIDEA. John B. Heppner & W. D. 
Duckworth 0 AO OTS 


E;XXPERIMENTAL HYBRIDIZATION BETWEEN PHYCIODES THAROS 
AND P. PHAON (NYMPHALIDAE). Charles G. Oliver __.___ 


NOTES ON THE BIOLOGY OF ZEGRIS EUPHEME (PIERIDAE). 
Steven Courtney 0 


HOSTPLANT RECORDS AND DESCRIPTIONS OF JUVENILE STAGES 


FOR Two RARE SPECIES OF EUVEIDES (NYMPHALIDAE). 


James L. B. Mallet & John T. Longino) 


FOSSIL LEAF-MINES OF BUCCULATRIX (LYONETIIDAE) ON ZELKOVA 
(ULMACEAE) FROM FLORISSANT, COLORADO. Paul A. 
Opler een a 


LIZARD PREDATION ON TROPICAL BUTTERFLIES. Paul R. Ehr- 

lich Anne H. Ehrlich Oo 
GENERAL NOTES 

Distinctiveness of Megisto c. cymela and M. c. viola (Satyridae). Charles 

Ge Olioer ye BG ee I PSO RSG NU err 


Long mating flights by male Hyalophora cecropia (L.) (Saturnidae). 
G. P. Waldbauer & J: G. Sternburg 200) So ee 


Hindtibial defensive spurs in the neotropical sphinx moth Amplypterus 
gannascus?’ Allen’ M. Young 20i\020 jee) GES oe 


Two species of skippers collected at antifreeze-filled pitfall traps in Ari- 
zona. Duane J. Flynn't Mogens C! Nielsen 2:0 a eee 


A one-fourth gynandromorph of Agriades rustica rustica (Edwards) from 
Wyoming (Lycaenidae). , Russell Rahn) 20 


Change in status of Catocala andromache race “benjamini’ (Noctuidae). 
ASE BROWER EOI tla Sige SLU) ee 


Mather S22 Bile 8 TOOT IRIN SUTURE AGI 


New hostplant records for Agonopterix clemensella (Oecophoridae). M. 
Berenbatini en i NS SI UCN SMES is GU hRA ee 


65 


76 
83 


87 


112 


119 
121 


132 


136 


145 


148 


153 
154 
155 


157 


158 


» 15y 


159 


160 


Volume 36 1982 Number 3 


ISSN 0024-0966 


JOURNAL 


‘ of the 

a 

r 9 

_ LEPIDOPTERISTS’ SOCIETY 

Published quarterly by THE LEPIDOPTERISTS’ SOCIETY 

F Publié par LA SOCIETE DES LEPIDOPTERISTES 
Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN 

i Publicado por LA SOCIEDAD DE LOS LEPIDOPTERISTAS 


14 December 1982 


THE LEPIDOPTERISTS’ SOCIETY 


EXECUTIVE COUNCIL 


CHARLES V. COVELL, JR., President LINCOLN P. BROWER, 
CLIFFORD D. FERRIS, Vice President Immediate Past President 
ALBERTO DfAz FRANCES, Vice President JULIAN P. DONAHUE, Secretary 
CLAUDE LEMAIRE, Vice President RONALD LEUSCHNER, Treasurer 


Members at large: 


R. L. LANGSTON K. S. BROWN, JR. F. S. CHEW 
R. M. PYLE T. C. EMMEL G. J. HARJES 
A. M. SHAPIRO E. H. METZLER 


The object of the Lepidopterists’ Society, which was formed in May, 1947 and for- 
mally constituted in December, 1950, is “to promote the science of lepidopterology in 
all its branches, .... to issue a periodical and other publications on Lepidoptera, to facil- 
itate the exchange of specimens and ideas by both the professional worker and the 
amateur in the field; to secure cooperation in all measures’ directed towards these aims. 

Membership in the Society is open to all persons interested in the study of Lepi- 
doptera. All members receive the Journal and the News of the Lepidopterists Society. 
Institutions may subscribe to the Journal but may not become members. Prospective 
members should send to the Treasurer full dues for the current year, together with their 
full name, address, and special lepidopterological interests. In alternate years a list of 
members of the Society is issued, with addresses and special interests. There are four 
numbers in each volume of the Journal, scheduled for February, May, August and 
November, and six numbers of the News each year. 


Active members—annual! dues $18.00 
Student members—annual dues $13.00 
Sustaining members—annual dues $25.00 
Life members—single sum $250.00 
Institutional subscriptions—annual $25.00 


Send remittances, payable to The Lepidopterists' Society, and address Samay. to: 
Ronald Leuschner, 1900 John St., Manhattan Beach, California 90266 U.S.A. 


Back issues of the Journal of the Lepidopterists’ Society, the Commemorative Vol- 
ume, and recent issues of the NEWS are available from the Treasurer. The Journal is 
$13 per volume, the Commemorative Volume, $6; and the NEWS, $.25 per issue. 


Order: Mail to Ronald Leuschner, 1900 John St., Manhattan Beach, California 90266 
U.S.A. 


Journal of the Lepidopterists’ Society (ISSN 0024-0966) is published quarterly by the 
Lepidopterists’ Society, a non-profit, scientific organization. The known office of publi- 
cation is 1041 New Hampshire St., Lawrence, Kansas 66044. Second class ec paid 
at Lawrence, Kansas, U.S.A. 66044. 


Cover illustration: Mature Larva of Eumorpha fasciata Sulzer (Sphingidae) feeding 
on Ludwigia sp. (Jussiaea) in southern Florida, where this hawk moth is generally found 
throughout the year. Original drawing by Mr. John V. Calhoun, 382 Tradewind Ct., 
Westerville, Ohio 48081, USA. 


JOURNAL OF 


Tue LeEpPIDOPTERISTS’ SOCIETY 


Volume 36 1982 Number 3 


Journal of the Lepidopterists’ Society 
36(3), 1982, 161-173 


SOIL- AND PUDDLE-VISITING HABITS OF MOTHS’ 


PETER H. ADLER 


The Frost Entomological Museum, Department of Entomology, 
The Pennsylvania State University, University Park, Pennsylvania 16802 


ABSTRACT. Ninety-three species of moths representing ten families were record- 
ed probing at soil and mud puddles over a four year period in central Pennsylvania. 
Nearly 99% of the 3417 individuals observed were males. Observations of Gracillari- 
idae and Lyonetiidae (97% males) are the first records for these families at soil. The 
natural history of the soil-visiting habits is described. Special mention is given those 
species of Geometridae and Notodontidae that pass large volumes of water through 
their gut as they drink from very wet substrates. Evolution of the soil-visiting habits 
and their relationship to animal excreta are discussed. 


The feeding habits of adult Lepidoptera are extremely diverse yet 
poorly understood. Gilbert and Singer (1975) stated that adults are 
more opportunistic and less specific in diet than larvae. The list of 
adult lepidopteran dietary sources other than floral nectar or extra- 
floral nectar (Downes, 1968) is extensive. Such sources include mud 
puddles, soil, and dung (Norris, 1936; Bauer, 1953; Sevastopulo, 1959, 
1974; Downes, 1973); urine (Owen, 1971); crushed bodies of conspe- 
cifics (Reinthal, 1966); moist campfire ashes (Howe, 1975); carrion 
(Reed, 1958; Payne & King, 1969; Shields, 1972; Downes, 1973; Niel- 
sen, 1977); saliva (refs. in Norris, 1936); exposed heads of basking 
turtles (D. L. Pearson, pers. comm.); soap suds (Farrell, 1979); lach- 
rymal secretions and pus (Banziger & Buttiker, 1969; Banziger, 1972): 
perspiration (Collenette & Talbot, 1928); plain salt (Skertchly, 1889); 
blood (Banziger, 1971, 1975); frog-hopper larval secretion (Lane, 1960): 
aphid honeydew (Manson, 1931); nectar gland secretion of lycaenid 
larvae (Gilbert, 1976); rotten fruit (Young, 1972); sound fruit (Har- 
greaves, 1936); cocoa seeds (Young, 1979); rotten seeds (Frost, p. 188, 


1 Accepted for publication as Paper No. 6406 on 17 March 1981 in Journal Series of the Pennsylvania Agricultural 
Experiment Station, University Park, Pennsylvania 16802, U.S.A. This paper is a contribution from the Agricultural 
Experiment Station Project No. 2070, “Biosystematics and Faunistic Survey of Insects.” 


162 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


1959); fermented milk (Howe, 1975); rotten cheese (Norris, 1936); 
borage plants (Pliske, 1975); tree sap (Tutt, 1897; Scott, 1973); red 
wine (Gomez, 1977); ink (Poulton, 1913); honey (d’Herculais, 1916); 
and pollen steeped in nectar (Gilbert, 1972). 

The present study surveys the soil- and puddle-visiting habits of 
moths in central Pennsylvania and presents notes on related feeding 
habits. Among most families of butterflies this behavior, usually re- 
ferred to as puddling, is extensively documented. However, only a 
few papers (Fassnidge, 1924; Collenette, 1934; Downes, 1973) indi- 
cate that moths, and particularly the nocturnal contingent, also visit 
soil and puddles. Norris (1936) and Downes (1973) summarize the 
families of Lepidoptera in which soil visitation occurs. 

In nearly all cases the Lepidoptera that do visit soil, puddles, car- 
rion, and animal excreta are predominantly males (review in Norris, 
1936). For example, Collenette (1934) found only 3% females among 
moths at damp sand in Brazil, and Downes (1973) observed 4.4% 
females among moths and butterflies at mud puddles in Ontario. One 
notable exception involves noctuids at perspiration in Brazil, where 
females represented 31% (Collenette, 1934). Females found at soil 
are usually old and wom (Clark, 1932). 

Arms et al. (1974) showed that the sodium ion stimulated puddling 
behavior in males of Papilio glaucus Linnaeus, and established that 
amino acids obtained from soil are incorporated into body proteins. 
Recently, Adler and Pearson (1982) demonstrated a difference in the 
total body sodium levels of male and female Pieris rapae Linnaeus, 
a species in which they likewise found a positive response to sodium. 

Lepidoptera that feed on dry substrates first moisten it with a bead 
of saliva passed down the proboscis and then reimbibe the solubilized 
nutrients in the manner described by Downes (1973). On the other 
hand, individuals that drink from wet substrates may discharge water 
droplets from the tip of the abdomen as they drink. 

As early as 1883, Dukinfield-Jones recorded that the Brazilian geo- 
metrid, Panthera apardalaria Walker, drank from wet stones in a 
stream for three hours and passed about 200 times its volume in liq- 
uid. Guppy (1952) observed Venusia cambrica Curtis drink from a 
bucket of water and pass the liquid in such a fashion as to resemble 
a “living siphon.” Clench (1957), in his observations of the geometrid, 
Dyspteris abortivaria Herrich-Schaeffer, and the drepanid, Drepana 
arcuata Walker, referred to the activity as “pumping.” | 

The discharging of liquid while at wet soil and puddles has also 
been observed in the Pyralidae (Welling, 1958); Hesperiidae (Roever, 
1964); Papilionidae (e.g., Reinthal, 1963; Welling, 1958; Jobe, 1977); 
Pieridae (Layard, 1883); and Lycaenidae (Tutt, 1897; pers. obs. of 


VOLUME 36, NUMBER 3 163 


TABLE 1. Summary of moths that visited soil and puddles during the summers of 
1977-1980. Species indicated by D consistently discharged droplets of liquid while 
drinking. Those marked I infrequently discharged. All others were not observed to 
discharge. Species are arranged alphabetically within families. 


Dis- 
charging 
Males Females behavior 


Gracillariidae 
Agrocercops perhaps striginifinitella (Clem.) 1 
Caloptilia cornusella Ely. 
C. perhaps stigmatella (F.) 
Leucospilapteryx venustella (Clem.) 
Parornix sp. 


WO & b OL 
| 


Lyonetiidae 
Bucculatrix sp. — il 


Gelechiidae* 
Genus, species unknown 


Tortricidae (including Olethreutidae) 


Ancylis metamelana (Wlk.) 

Cenopis reticulatana Clem. 

Grapholita eclipsana Zeller (diurnal) 
Olethreutes albiciliana (Fernald) (diurnal) 


Pyralidae 


Anania funebris glomeralis (Wlk.) (diurnal) 
Argyria nivalis (Dru.) 

Blepharomastix ranalis (Gn.) 

Crambus turbatellus WIlk. 

Desmia funeralis (Hbn.) 

Mutuuraia mysippusalia (Wlk.) 

Ostrinia nubialis (Hbn.) 

Pyrausta pertextalis Lederer 

Sylepta fluctuosalis (Lederer) 

Pyralid sp. 


_— 


coer 
| 


— 


= 
i) bo 
RePWr WON Fe WO 


Pterophoridae 


Oidaematophorus homodactylus (Wlk.) 
O. monodactylus (Linn.) 


me 
| 


Drepanidae 
Eudeilinea herminiata (Gn.) 4 = 


Geometridae 


Anacamptodes ephyraria (Wlk.) 
Anagoga occiduaria (WIlk.) 
Anavitrinella pampinaria (Gn.) 
Antepione thisoaria (Gn.) 

Biston betularia cognataria (Gn.) 
Cepphis armataria (H.-S.) 
Chlorochlamys chloroleucaria (Gn.) 
Coryphista meadii (Pack.) 
Cyclophora myrtaria (Gn.) 

C. packardi (Prout) 

Dyspteris abortivaria H.-S. 672 


— 
| eto SG Gui as 
| 


| 00 | 


164 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


TABLE 1. Continued. 


Dis- 
charging 
‘Males Females _ behavior 
Dystroma hersiliata (Gn.) ] real 
Ectropis crepuscularia (D. & S.) 1 — 
Euchlaena irraria (B. & McD.) rif = 
Euphyia unangulata intermediata (Gn.) 14 — I 
Eupithecia sp. 1 — 
Eusarca confusaria Hbn. 69 — 
Haematopis grataria (Fab.) 1 = 
Heliomata cycladata Grote (diurnal) 2) — 
H. infulata Grote (diurnal) ] — 
Heterophleps triguttaria H.-S. 44 = D 
Homochlodes fritillaria (Gn.) — 1 
Hydrelia lucata (Gn.) 11 = D 
Hydria prunivorata (Ferguson) 90 — I 
Hydriomena perfracta Swett. 1 — 
Idaea demissaria (Hbn.) 1 1 
Iridopsis larvaria (Gn.) 12 i 
Itame argillacearia (Pack.) 76 = 
I. pustularia (Gn.) 3 2 
Lobophora nivigerata Wlk. 28 a 
Lomographa semiclarata (Wlk.) (diurnal) 700 25 
Melanolophia canadaria (Gn.) 14 a 
M. signataria (Wlk.) 1 — 
Mesoleuca ruficillata (Gn.) 24 — 
Metanema determinata WIk. 36 — I 
M. inatomaria Gn. 33 — I 
Metarranthis angularia B. & McD. 10 — 
Nematocampa limbata (Haw.) 19 == D 
Nemoria bistriaria Hbn. 4 == 
N. rubrifrontaria (Pack.) 1 — 
Orthonama centrostrigaria (Wollaston) By at I 
Plagodis alcoolaria (Gn.) 16 — 
P. fervidaria (H.-S.) ile} — 
P. phlogosaria (Gn.) 33 — 
Probole alienaria H.-S. 72 — D 
P. amicaria (H.-S.) 91 — D 
Scopula inductata (Gn.) 18 — 
S. limboundata (Haw.) 85 2 
Semiothisa bisignata (Wlk.) = l 
Sicya macularia (Harr.) 5 — 
Synchlora aerata (F.) 14 — 
Xanthorhoe ferrugata (Clerck) 58 — I 
X. lacustrata (Gn.) 15 — 
Xanthotype urticaria Swett. 118 — I 
Notodontidae 
Clostera albosigma Fitch 210 — D 
Gluphisia septentrionis Wlk. DH — D 
Noctuidae 
Bomolocha baltimoralis Hbn. 13) — 


Enargia decolor W\k. 3 — 


VOLUME 36, NUMBER 3 165 


TABLE 1. Continued. 


Dis- 
charging 
Males Females behavior 


Hypena humuli Harr. 
Hyperstrotia sp. 
Lithachodia carneola (Gn.) 
Orthosia sp. 

Palthis angulalis Hbn. 
Psychomorpha epimenis Dru. (diurnal) 
Renia discoloralis Gn. 

R. factiosalis (Wlk.) 

R. sp. 

Tarachidia erastrioides (Gn.) 
Zale undularis (Dru.) 


Sh eres bapereer sr cel 


* R. L. Mangan (pers. comm.) commonly observed the pink bollworm, Pectinophora gossypiella (Saunders) feed- 
ing from soil in irrigated cotton fields of Arizona on very warm days. 


Celastrina argiolus pseduoargiolus (Boisduval & LeConte)). Banzig- 
er (1972) and Reid (1954) provide examples of nocturnal Lepidoptera 
discharging drops of liquid while feeding on the eye secretions of 
various animals. Certain hesperiids turn their abdomens anteroven- 
trally and expel a drop of liquid in order to moisten a substrate and 
subsequently imbibe the drop (refs. in Norris, 1936; Hessel, 1966; 
Jobe, 1977; pers. obs. of Erynnis juvenalis Fabricius and E. baptisiae 
(Forbes)). 


STUDY AREA AND METHODS 


I observed nocturnal Lepidoptera with the aid of a head lamp from 
June through the third week of August each year from 1977 to 1980 
(2130-0130 EDST). All field work was conducted in the Scotia Bar- 
rens of Centre County, Pennsylvania (approximately 5.7 km west of 
State College) along a 2.4 km stretch of gravel-dirt road leading south 
of Ten Acre Pond. The soil of the Scotia Barrens is characterized as 
Morrison loamy-sand. 

Only those individuals with their proboscis extended to the sub- 
strate were counted. Individuals were sexed in the field and several 
representatives of each species were collected for identification. 


RESULTS 


During the four years of observation encompassed by this study I 
recorded 3417 moths, representing 93 species in ten families, at damp 
soil, puddles, ponds and their edges (Table 1). Observations of Gra- 
cillariidae and Lyonetiidae at soil are new family records. Females 
probing at damp soil comprised 1.3% of all individuals and repre- 


166 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


sented only 16 species. Notably, half of these species were unrepre- 
sented by males. 

All females in Table 1, with the exception of Zale undularis Drury, 
were old and wor; whereas, many of the males appeared fresh. How- 
ever, senescent males of some species were common at damp soil. 
For example, 40% of Eusarca confusaria Hubner males (n = 63) were 
fresh (less than two days old), as determined by the wing condition 
of marked-recaptured individuals, while 31% were middle-aged and 
29% were old (more than seven days old). A mark-recapture study of 
E. confusaria revealed that the same male may visit soil on more than 
one night. 

Although flowers (Asclepias syriaca L., Melilotus alba Desr., Achil- 
lea millefolium L., Spiraea latifolia (Ait.) Borkh., Solidago spp., and 
Eupatorium spp.) bloomed plentifully in the study area, soil visitation 
was far more frequent than nocturnal flower visitation among males 
of all species in Table 1, except E. confusaria. I observed 303 E. 
confusaria males feeding during July 1977 but only 19.8% of these at 
damp soil. All 171 E. confusaria females found feeding were at flow- 
ers. 

Some areas of the soil visited by moths were intimately associated 
with the recent presence of animals. Nocturnal Lepidoptera fed on 
bird droppings (males of Heterocampa manteo Doubleday and E. 
confusaria), dead frogs and rabbits, animal urine, and in the footprints 
of deer. (Although five males of Prochoerodes transversata (Drury) 
fed on the exudates of a dead rabbit, they were never observed at 
soil.) Aggregates of nocturnal moths were sometimes associated with 
animal-related substrates and restricted patches of moisture. 

I often found nocturnal moths probing on soil-related substrates 
such as the damp walls, floors, and ceilings of crumbling cement struc- 
tures in the abandoned (since the 1920's) community of Scotia. Most 
species found on the cement also visited soil with the exception of 
two female Catocala ultronia Hitbner, one female C. ilia Cramer, 
one female Euparthenos nubialis Hubner, and one female Amphi- 
pyra pyramiodoides (Guenée). Although the observations of the latter 
four species are unique, it is of note that these species often rested 
on the cement structures during the day. 

The species that visited soil and puddles could generally be char- 
acterized as those that passed droplets of liquid as they drank (Figs. 
1 and 2), those that did so infrequently (less than 10% of the individ- 
uals of a given species), or those that did not (Table 1). Species that 
habitually discharged invariably drank from puddles or soil with a 
thin film of water. Non-discharging and infrequently discharging 
species preferred damp soil without standing water and sometimes 


VOLUME 36, NUMBER 3 167 


Fic. 1. Male Dyspteris abortivaria Herrich-Schaeffer, a typical discharging species, 
at wet soil. Arrow indicates formation of a droplet of liquid at the tip of the abdomen. 


drank from the cracked bottoms of drying puddles. I observed no 
female of any species discharge while at soil. 

The position in which moths held their wings while at soil or pud- 
dles was characteristic for each species. Among geometrids, approx- 
imately 80% including all discharging species held their wings toward 
the perpendicular (Fig. 3). Other geometrids such as Hydria pruni- 
vorata (Ferguson), Lobophora nivigerata Walker, both Metanema spp., 
and Scopula limboundata (Haworth) held their wings against the sub- 
strate. 

Individuals that settled at soil or puddles generally remained sta- 
tionary for several hours. The formation of liquid droplets at the tip 
of the abdomen and/or antennal palpation was often the only visible 
activity. All individuals were seemingly unaffected by light or close 
proximity of the observer. When moths completed their drinking or 
were touched they were capable of immediate flight except Gluphisia 
septentrionis Walker and Clostera albosigma Fitch, which vigorously 
vibrated their wings before actual flight. 

G. septentrionis and C. albosigma restricted their drinking to areas 
of soil associated with a puddle or pond. C. albosigma often drank 
from scum and algal mats that floated on the surface, as well as from 
the edges of the water. This moth occasionally floated on shallow 
water while drinking. G. septentrionis frequented pond and puddle 
edges, where I observed individuals drink for more than an hour on 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Fic. 2. Male Gluphisia septentrionis Walker discharging a jet of water (indicated 
by arrow). Above, substrate is a wet board. Below, rifle (.22 caliber) shell case indicates 
size. 


wet boards, stones, moss, or head-down on the stems of emergent 
vegetation. All observations indicated that these species discharged 
water from the tip of the abdomen at a very rapid rate. G. septentrionis 
forcibly expelled rhythmic jets that traveled distances up to 30 cm 
(Fig. 2) whereas C. albosigma discharged large drops that diffused into 
the surrounding water. Among the geometrids that discharged liquid, 


VOLUME 36, NUMBER 3 169 


Fic. 3. Male Dyspteris abortivaria Herrich-Schaeffer showing usual position of the 
wings while visiting soil. Only the apical portion of the proboscis is applied to the 
water film. 


drop size was generally proportional to body size, and when water 
was freely available, drops were produced at the rate of one or two 
per second. Suspended soil particles were also passed with the water. 
No other species at this site passed water through its system as 
rapidly and in such quantities as G. septentrionis. A typical individual 
on a wet cinder road discharged 5.7 ml in 40 minutes (15 jets/min) 
along with 10.0 mg (dry weight) of particulate. An individual brought 
into the lab and offered a 10 uM solution of NaCl passed 22.7 ml in 
78 minutes (35 jets/min or 510 times its wet weight in liquid). Bro- 
mophenol red placed in the drinking water of three moths in the field 
was passed with the first ejection, i.e., within three to five seconds. 


DISCUSSION AND CONCLUSIONS 


The preceding results emphasize the highly developed habit of soil 
and puddle visitation in the ditrysian Lepidoptera, while a synopsis 
of the literature (Norris, 1936) reveals the widespread geographical 
nature of the habit. This attraction to soil and puddles probably evolved 
from water-drinking, a behavior necessary for maximum fitness in most 
Lepidoptera (Norris, 1934, 1936). The drinking behavior may have 
acquired further significance as individuals that satisfied their water 
requirements at soil or puddles accrued additional benefits, despite 
predation risks (Morris, 1953; pers. obs. of Chipping, Song and White- 


170 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


crowned Sparrows and Eastern Towhees on Lomographa semiclarata 
(Walker) and Erynnis juvenalis and Common Grackles on Pieris ra- 
pae). 

Common (1975) believes that the simple haustellate mouthparts of 
Lepidoptera prefaced the appearance of Angiosperms and further notes 
that the evolution of a functional proboscis would have allowed the 
early Lepidoptera with mandibles to move into drier places if they 
imbibed moisture droplets. If, in fact, the soil-visiting habits evolved 
from the habit of water-drinking, one would expect the more primitive 
Lepidoptera with a functional proboscis to visit soil. 

Until the previously unrecorded observations of the Gracillariidae 
and Lyonetiidae (Table 1), soil visitation was believed generally re- 
stricted to the higher families of Lepidoptera. With the addition of 
these records, the habit assumes significance as a more widespread 
behavioral and physiological character. As in the higher Lepidoptera, 
a paucity of female soil visitors is evident among the Gracillariidae 
and Lyonetiidae, and may be considered characteristic of the soil- 
visiting Ditrysia. 

The results of this survey increase the numbers of Geometridae 
that are known to characteristically discharge droplets of liquid while 
drinking from soil and puddles and provide the first records of this 
behavior for the Notodontidae. Drinking at dry or damp soil and drink- 
ing at open puddles appear to be variants of the same behavior. Whether 
or not a discharge occurs is related to the choice of microhabitat (often 
species-specific), which is, in turn, a function of water availability. 
This behavior is quite similar to that of aphids which pass copious 
amounts of plant juices in order to extract nutrients present in trace 
quantities (Mittler, 1958). I have also seen scores of Halysidota tes- 
sellaris (Smith) imbibe superfluous volumes of dilute Asclepias nectar 
just after a rain and discharge large droplets of liquid but have not 
observed this situation on nights without rain. 

An anatomical and histological study of the alimentary tract of G. 
septentrionis, including a comparative study of the sexes, with respect 
to the rapid and forceful passage of large quantities of liquid would 
be informative. My preliminary dissections suggest that, superficially, 
the gut of the male does not differ markedly from that of an actively 
feeding noctuid, Plusia gamma Linnaeus (Mortimer, 1965). 

All feeding habits associated with animal excreta probably evolved 
from the habit of visiting soil, and Downes (1973) is probably correct 
in assuming that animal excreta provide higher levels of attraction 
than soil and puddles. Sodium represents one common factor linking 
soil and puddles with animal excreta as nutrient substrates and has 
been suggested (Poulton, 1917) and implicated (Arms et al., 1974; 


VOLUME 36, NUMBER 3 ey 


Adler & Pearson, 1982) as the feeding stimulus. However, the exclu- 
sively lachryphagous noctuid, Lobocraspis griseifusa Hampson, serves 
as a caveat that not all lepidopteran feeding habits associated with 
animal excreta involve uptake of similar nutrients. The fact that both 
sexes are commonly involved (Banziger, 1975), and the species is 
unique among Lepidoptera thus far studied in producing pro- 
teinases (Banziger, 1972), sets it apart from typical soil and animal 
associated Lepidoptera. 

Viewed as a whole, there seems little question that soil and puddle 
visitation represent an integral part of the biology of many taxonom- 
ically diverse Lepidoptera. Species such as G. septentrionis that dis- 
charge large volumes of liquid as they drink afford excellent subjects 
for quantifying the dietary aspects of soil and puddle visitation through 
determination of substances in the imbibed medium versus the dis- 
charge. 


ACKNOWLEDGMENTS 


I thank Mr. F. D. Fee, Dr. K. C. Kim, Dr. D. L. Pearson, and Dr. C. W. Pitts, The 
Pennsylvania State University, for their helpful comments on the manuscript. I also 
thank Dr. R. L. Brown, Mississippi State University, for his determinations of the 
diurnal Tortricidae; Dr. D. Davis, Department of Entomology, Smithsonian Institution, 
for his determinations of the Gracillariidae and Lyonetiidae; and Dr. R. W. Hodges, 
Systematic Entomology Laboratory, United States Department of Agriculture, for his 
determination of the Gelechiidae. I especially thank Dr. D. C. Ferguson, Systematic 
Entomology Laboratory, U.S.D.A., for his kind assistance and determinations of all 
other Lepidoptera in this study. I also extend my gratitude to the many individuals, 
especially C. S. Brooks, who accompanied me into the field. 


LITERATURE CITED 


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1975. Skin-piercing blood-sucking moths I: Ecological and ethological studies 

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VOLUME 36, NUMBER 3 173 


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Journal of the Lepidopterists’ Society 
36(3), 1982, 174-177 


REDUNDANCY IN PIERID POLYPHENISMS: PUPAL 
CHILLING INDUCES VERNAL PHENOTYPE IN 
PIERIS OCCIDENTALIS (PIERIDAE) 


ARTHUR M. SHAPIRO 


Department of Zoology, University of California, 
Davis, California 95616 


ABSTRACT. Chilling young pupae of Sierra Nevada Pieris occidentalis induces 
the vernal-alpine “calyce” phenotype as effectively as rearing on short days. Temper- 
atures of 6°C or less sustained for 10 days or more appear equally efficacious. This 
redundancy in phenotypic-induction mechanisms parallels that found in various other 
butterflies, and is underlain by apparent genetic variation in sensitivity within popu- 
lations. 


The Western white, Pieris occidentalis Reakirt, has two seasonal 
phenotypes once considered separate species (Edwards, 1876). Sha- 
piro (1973) demonstrated that rearing on a short day (10L:14D) with- 
out chilling induced the heavily-marked phenotype “calyce” in Bo- 
real Ridge, California stock, and that temperatures of 10°C were 
ineffective in doing so under long-day (14L:10D) conditions. Later, 
Shapiro (1978) demonstrated that redundancy existed in the pheno- 
typic-induction systems of various multivoltine Pieridae, including 
the closely-related species P. protodice Boisduval and LeConte. This 
discovery prompted a re-evaluation of thermal influences in pheno- 
typic determination in P. occidentalis; specifically, chilling of long- 
day pupae at lower temperatures than used heretofore. 

Ova were obtained from a single wild female collected in Donner 
Pass, of the Sierra Nevada, Nevada County, California (2100 m), 7 
August 1980. This locality is about 4.5-km from Boreal Ridge and at 
similar elevation. Rearing was done under our standard conditions 
(Shapiro, 1975) with continuous light at 25°C on Lepidium virginicum 
L. var. pubescens (Greene) Thellung (Cruciferae), a natural host at 
Donner Pass. Allocation of pupae to treatments was randomized to 
obviate effects of the sequence of oviposition on offspring quality. 
Control pupae were held at the rearing conditions. Experimental pu- 
pae were refrigerated as close to eight hours after pupation as possible 
(effectively +1 h) and held in one of four regimes: 6°C for 10 days; 
5°C for 14 days; 2°C for 14 days; 10 days at 6°C followed by 2 days at 
25°C followed by 7 days at 2°C. After chilling, the pupae were re- 
turned to 25°C and allowed to develop and eclose. Mortality was neg- 
ligible (4/154). Adults were classified into three phenotypic grades 
based on the degree of melanization of the ventral hindwing; stan- 
dards are shown in Fig. 1 and correspond to those used by Shapiro 


VOLUME 36, NUMBER 3 175 


Fic. 1. Phenotypic standards for Pieris occidentalis, ventral surfaces, males at left. 
Top row, light; middle, intermediate; bottom, dark. All bred ex Donner Pass. 


(1973). The results are given in Table 1. They were analyzed as fol- 
lows: a log-linear model (Fienberg, 1977) was fitted to the 5 x 3 con- 
tingency table for pooled sexes. Using the large-sample distribution 
of the natural logarithms of the frequencies, it was possible to express 
the subsequent tests in terms of linear contrasts. When the hypothesis 
that the treatments have no effect on the phenotypic distribution was 
tested (pooled treatments vs. controls), the resulting y” statistic was 


176 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 1. Phenotype distributions in a brood of 150 Pieris occidentalis from one 
Donner Pass, CA, female, reared on 24L/25°C, subjected to five different pupal tem- 
perature regimes. 


Males Females Both sexes 
Inter- Inter- Inter- 
Treatments: Dark mediate Light Dark mediate Light Dark mediate Light 
A. Control (unchilled) 4 12 15 1 10 Dil 5 22 36 
B. 10 days at 6°C 2 11 4 4 5 1 6 16 5 
C. 14 days at 5°C 4 6 0) ma 6 1 11 12 il 
ID), 14! claws ate 2G 3 6 0 4 6 0 7 12 0 
He lOidaysvat 62; 
then 2 days at 25°C, 
then 7 days at 2°C 3) 3 2 3 3 1 8 6 3 
> chilled 14 26 6 18 20 3 Oo 46 9 
Statistical comparisons (pooled sexes): A XB: x’; = 11.09 (0.01 < P < 0.025) 
IN S< (Ck x’, = 18.25 (P < 0.005); 
AxXD: x’, = 13.88 5 < Hour 
AXE: x’, = 13.64 (P < 0.005); 
B x E: x’, = 3.12 (0.1 < P); 
C x D: x’, = 0.59 (0.1 < P). 


33.89 (df = 8, P < 0.005). Comparisons between the control and the 
individual treatments were all significant (Table 1), while two sample 
comparisons were not. I therefore concluded that the phenotype of 
P. occidentalis is inducible by pupal temperature exposure but that 
it behaves as a threshold phenomenon with the switch occurring at 
some temperature between 10°C and 6°C. 

Though further refinement of the effects is likely using larger sam- 
ples and perhaps analyzing the sexes separately, it is evident that P. 
occidentalis adheres to the pattern of P. napi L., P. protodice, and 
Colias eurytheme Bdv. in having built-in redundancy in its pheno- 
typic-induction system, with short days irreversibly determining dark 
phenotype but, with long day—light phenotype decisions reversible 
by subsequent chilling. Also, the response is not all-or-none; some 
seemingly inappropriate phenotypes are produced in most groups, 
and the distribution of this phenomenon exceeds the variance in age 
at refrigeration. Thus differences in sensitivity to chilling, as to pho- 
toperiod, may be genetically determined. Extreme “calyce” pheno- 
types (Shapiro, 1978) probably represent the combination of larval 
short-day exposure and post-diapause pupal chilling. Since my 1973 
paper I have examined many populations of this complex from the 
Rocky Mountains, the Sierra Nevada, and Alaska, all of which retain 
both “calyce”’ and estival occidentalis phenotypes. 


VOLUME 36, NUMBER 3 ear, 


ACKNOWLEDGMENTS 


I thank Mike Miller for statistical advice, Marc Minno for technical assistance, and 
the Department of Zoology for support of field and lab studies in the Nearctic Pierinae. 


LITERATURE CITED 


EDWARDS, W. H. 1876. Catalogue of the diurnal Lepidoptera of America north of 
Mexico. Trans. Amer. Entomol. Soc. 6:1-68. 

FIENBERG, S. E. 1977. The analysis of cross-classified categorical data. M.I.T. Press, 
Cambridge, Mass. 151 pp. 

SHAPIRO, A. M. 1973. Photoperiodic control of seasonal polyphenism in Pieris occi- 
dentalis Reakirt (Lepidoptera: Pieridae). Wasmann J. Biol. 31:291-299. 

1975. Photoperiodic control of development and phenotype in a subarctic 

population of Pieris occidentalis (Lepidoptera: Pieridae). Canad. Entomol. 107: 

T15-179. 

1976. The biological status of Nearctic taxa in the Pieris protodice-occidentalis 

group (Pieridae). J. Lepid. Soc. 30:289-300. 

1978. The evolutionary significance of redundancy and variability in pheno- 

typic-induction mechanisms of Pierid butterflies (Lepidoptera). Psyche 85:275- 

283. 


Journal of the Lepidopterists’ Society 
36(3), 1982, 178-184 


ROOST RECRUITMENT AND RESOURCE UTILIZATION: 
OBSERVATIONS ON A HELICONIUS CHARITONIA L. 
ROOST IN MEXICO (NYMPHALIDAE) 


D. A. WALLER AND L. E. GILBERT 


Department of Zoology, University of Texas, 
Austin, Texas 78712 


ABSTRACT. <A communal roost of individually marked Heliconius charitonia L. 
(Nymphalidae) butterflies was observed over a three week period in southern Mexico. 
Pollen plants (Anguria, Cucurbitaceae), which serve as adult resources for these but- 
terflies, were monitored for butterfly visits during this time. Roost members were gen- 
erally using separate pollen plants from non-roost member H. charitonia. Predation on 
a roost member was observed at one of these plants. Fresh H. charitonia were observed 
associating with roost members first at a pollen plant during the day and later that 
evening at the roost site, indicating that the new recruits followed the older butterflies 
to the roost. Observations are discussed in regard to current hypotheses about gregar- 
ious roosting. 


Many species of birds, solitary bees and wasps, bats and butterflies 
roost communally. In most cases, members depart from the roost on 
a circadian schedule to forage, returning to the site to sleep. Roosts 
may be diurnal or nocturnal, seasonal or permanent, mono- or multi- 
specific. 

There are two major hypotheses concerning why animals roost to- 
gether. One hypothesis suggests that communal roosts protect mem- 
bers from predation, either because animals in groups are quickly 
alerted to predator presence (Gadgil, 1972), or because the species is 
unpalatable (Benson, 1971; Turner, 1975). The second hypothesis 
proposes that roosts serve as centers of information exchange about 
food resources (Ward & Zahavi, 1973; Gilbert, 1975). At present, there 
are too few data on roost dynamics in any species to fully assess the 
relative importances of these two hypotheses. 

Some species of the brightly-colored Neotropical Heliconius (Nym- 
phalidae) butterflies characteristically form nocturnal communal roosts. 
Members of these roosts often home repeatedly to the same site every 
night (Benson, 1971). Gilbert (1975) has suggested that new recruits 
to Heliconius roosts follow experienced roost members from the roost 
site when they forage in order to learn the locations of pollen plants, 
which serve as important adult food sources for these long-lived but- 
terflies (Dunlap-Pianka et al., 1977). Following behavior by conspe- 
cific Heliconius has often been observed in the field, but there is no 
information on following by members of the same roost. Similarly, ~ 
there is no substantial information on patterns of resource utilization 
by roost members of Heliconius butterflies. 


VOLUME 36, NUMBER 3 179 


In our study of a Heliconius charitonia L. roost in Mexico, we ob- 
served fresh butterflies associating with roost members first on a pol- 
len plant during the day, and later that evening at a roost site, indi- 
cating that the new recruits found the roost site by following the older 
butterflies. We also obtained evidence that roost membership is closely 
tied to resource use. 


METHODS 
Heliconius Butterflies 


Heliconius are aposematic Neotropical butterflies with limited home 
ranges (Ehrlich & Gilbert, 1973). Individuals often live and reproduce 
for six months or longer. Both sexes make repeated visits to adult food 
plants for pollen and nectar, and to larval host plants for ovipositions 
(females) and mate-finding (males). It is therefore possible to monitor 
home range movements and resource utilization by individual roost 
members. 

Heliconius charitonia is a brown and yellow zebra-striped inhab- 
itant of forest edge and secondary growth habitats. This species forms 
low, cryptic nocturnal roosts, sometimes with other Heliconius species. 


Study Site 


Observations were made at the Estacion de Biologia Tropical “Los 
Tuxtlas’” UNAM, near the town of Catemaco, Veracruz, Mexico. The 
Station is located on 700 ha of primary tropical rainforest and second- 
ary growth. Altitude ranges from 150 m to 530 m. The mean annual 
temperature is 24°C, and the average precipitation is 4560 mm per 
year. This study was conducted in July and August 1978, approxi- 
mately one month into the wet season. 


Roost 


The roost observed during this study was located next to a stream 
bed in a clearing that had been created by a tree fall (see Fig. 1). Two 
adjacent but discrete subroosts were used by the butterflies; these 
were approximately 3 and 4 meters high, respectively. Butterflies oc- 
casionally roosted high in branches above both subroosts. 


Adult Food Plants 


Anguria tabascensis Donn. Smith (Cucurbitaceae) lianas were the 
major pollen sources for H. charitonia at the Station. Eight of these 
plants were monitored for visits by Heliconius butterflies. Plants Al, 
A2, A3, A4, A5, A7 and A8 were flowering at eye level, and numbers 
of butterfly visitors could be read off the wings without disturbing 


180 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Fic. 1. Map of roost and surrounding area at the Estacion Biologia Tropical “Los 
Tuxtlas’”” UNAM, Veracruz, Mexico, including locations of Anguria pollen plants A1, 
A2, A3, A4, A5, A6, A7 and A8. Roost member H. charitonia captures and sightings 


VOLUME 36, NUMBER 3 181 


the butterflies. A6 was a large vine flowering 22 meters in the canopy. 
Butterfly visitors to A6 were identified using binoculars and a Questar 
3 telescope. Fig. 1 shows the locations of the roost and of Anguria 
plants Al—A8. 


Procedure for Observation 


Butterflies were observed on the roost through binoculars mornings 
and evenings from 18 July—6 August 1978, and evenings only from 6 
August-10 August 1978. Roost members were caught at Anguria plants 
during the day, or netted as they left the roost in the morning. They 
were numbered at these times on the forewings with black marking 
pen. Butterflies were scored for sex and wingwear as fresh (F: less 
than 1 month old), intermediate (I: between 1 and 3 months old), or 
worn (W: over 3 months old), following Ehrlich and Gilbert (1973). 


RESULTS 
Roost Fidelity 


When the roost was first discovered on 17 July 1978, it was com- 
posed entirely of wom and intermediate butterflies. No fresh butter- 
flies were observed at the roost site until ten days later on 27 July. 

Individual butterflies were highly faithful to the roost (see Fig. 2). 
The average roost member spent 11.4 nights on the roost during the 
three week period. Of those butterflies that spent more than one night 
on the roost (24 out of 27 butterflies), the average roost member was 
on the roost 87% of the nights during the time the butterfly was first 
and last observed at the site. Other studies of H. charitonia roosts 
have found roost membership to be less constant (Young & Carolan, 
1976; Young, 1978). We attribute some of the high fidelity we ob- 
served to our avoidance of disturbing roosting animals for marking. It 
has been established that capture may drastically affect recapture of 
mud-puddling butterflies (Singer & Wedlake, 1981). 


Resource Utilization 


Another factor that may have contributed to the high roost fidelity 
observed in this study was the presence of a large pollen source 50 
m from the roost site. This Anguria plant, A6, was located 22 m in 


— 


were confined within the dotted line surrounding the roost site. Non-roost member H. 
charitonia captures and sightings are indicated by x’s. Dark solid lines represent roads 
and paired wavy lines represent streams. (We thank Alejandro Estrada for access to 
this map which was made by his group for howler monkey population studies.) 


182 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


July August 

i (SC Mw 19 2O Bl 22 2 BA 2 2 27 2 2. 30 Bi i 2 3 4 5) 6 7 8 9 10 1) 

MEMEMEMEMEMEMEMEMEMEMEMEMEMEMEMEMEMEMEME 13 E E E E 
A Wl Wheelie -SReserseaSaass= 
7 
10 F VWtat ----- wae een nnn 5 55a n= eee ann 2H = 4 --==---- 
12M Fo weeaeaaaa-a=-  ------=-- a Seana sees aaa wea wee 
14M IW --- - = ae nnn nn 
ISM F 2 ssecs So SSS 
DA VN A = eS SSS SSS SSS SS Se SSS 
24 F IWtat 
26 F I Se Sr sn nessa =a === == 
28M W m-- 00 = =-------------- = -oo-- = = eee nnn =- 
30M WO nnn nn =a =H === = naoa- - 0 =oa=- = seseenaa==== 
32M I ma= 0 anna ana- 0 = 2 = === =---=-- setae nana nna -==- = ween anna 
34 F W ea- 0 2a 25 ---=-==----=- = Je ip A Peet Ver) 253 = 
36M Wo tne = 
38M IWtat 4 4242 242 22 annem = seen anne 
AQ Mh ea --- 
42M IW Rann sa 8a 
44 M Wtat Soo igo 
46 M Wtat carne 
ABM WR) 0 SESE SSSR SSeS SSSR = SS ~ enna nn 
SOMRA IE ok) peaodeob | lle ions Sateen a ui at BS = seces = 
G0) 1) = 
62 M F a 
C6 ye Pe DEED Ee Oe eR cote Roe et a ees oe ce ea a ae fe. 
CBM ORS EA Ba a ie Bi RE ie te RS Tae aca lg: cas a a le aaa 
72 i We Fy ¥ 
U3 ie + 
Fresh unmarked butterflies 2i 3s 333 3 VN i TW ] 1 2 1 ] ] 2 


6 7 3 OS AWA A AWA} AT BST Oe 2 Seale 4 
Total of Marked Butterflies on Roost 
7 8 hy VA 1 1 iG) Ne 7 16 16 1S We We NB WS NG WH 1 WA WA je WA 3 4 
Total of Marked Roost Members Known Alive 


Fic. 2. Mornings (M) and evenings (E) butterflies were observed on the roost. Sex 
(S) and condition (C) of roost members are indicated. 


the canopy and bore more than 33 male inflorescences. More than 
half of the roost members (11 of 19 marked butterflies at the time of 
observation) were seen visiting flowers at A6, often repeatedly. A6 
was observed for a total of six hours over three separate days. 

Other butterfly species, including other heliconiines, also visited 
A6. However, with the exception of one worn, unmarked butterfly, 
the H. charitonia observed at A6 were all roost members. The almost 
exclusive use of this resource by roost members was especially strik- 
ing since non-roost member H. charitonia were repeatedly observed 
flying in the road below A6. 

Roost members also visited two Anguria that were growing at the 
edge of a cultivated field just north of the roost site. In four hours of 
observations (three separate days), seven roost members (Numbers 7, 
10, 12, 15, 24, 26, and 42) were seen at A7. Only one non-roost mem- 
ber was caught and marked at A7. In over three hours of observations 
(four separate days), two H. charitonia roost members (Numbers 24 
and 58) made repeated visits to A8. Most of the roost members at A7 
were also seen in the canopy at A6 (Numbers 7, 10, 12, 26, and 42), 
but visitors to A8 were never seen at A6. 

Anguria plants Al, A2, A3, A4 and A5 were monitored for Helico- 
nius visits throughout the study. Heliconius erato, H. doris, H. isme- 
nius, and H. charitonia were caught and marked at these plants from 
7 July 1978, through 7 August 1978. No roost members were ever 


VOLUME 36, NUMBER 3 183 


observed to visit these flowers, and none of the 21 H. charitonia 
marked at these plants appeared at the roost site. 


Predation at Resource 


One worn roost member (Number 46, W male) was killed but ap- 
parently not eaten in the presence of roost mates at A6 by a tanager 
(Thraupidae). This observation made with the Questar suggests that, 
through their use of the same favorite pollen plants, roost mates pro- 
vide a context for the operation of visual selection by predators, even 
away from the roost. This is in accord with Benson’s (1971) and Turn- 
er s (1975) models that link roosting behavior with distastefulness and 
aposematic coloration in Heliconius butterflies. 


Roost Recruitment 


On the morning of 27 July 1978, two fresh H. charitonia butterflies 
appeared at A6 in the canopy, one with a distinct reddish cast to its 
wings. These fresh butterflies associated with roost members No. 38 (I 
male) and No. 40 (W male) on the flowers. That evening, two fresh but- 
terflies, one with a distinct reddish cast to its wings, appeared at the 
roost site for the first time since the commencement of observations 
10 days previously. One new recruit roosted with the group, and the 
other roosted with No. 38 on a branch away from the other butterflies. 
Although it cannot be documented that the two fresh butterflies sight- 
ed at A6 were the same that joined the roost in the evening, the 
circumstantial evidence strongly suggests that they were. 


DISCUSSION 


Our observations indicate that roost members were using separate 
pollen sources from those of non-roost member H. charitonia. Wheth- 
er or not roost members learn the locations of these plants from each 
other remains to be investigated. This finding does suggest, however, 
that roost membership is somehow tied to patterns of adult resource 
utilization. If roost members associate in space and time at pollen 
plants, then predators need not visit the roost to learn or reinforce 
avoidance of one roost member through experience with another. 

New recruits associated with roost members first on a pollen plant, 
and later at the roost site, apparently following the experienced but- 
terflies to the roost. This observation suggests a mechanism for roost 
recruitment. Mature butterflies continually canvass larval host plants 
(Passiflora spp., Passifloraceae), with females in search of oviposition 
sites and males in search of female pupae. New butterflies emerging 
in the vicinity of Passiflora plants may be attracted to older butterflies 
when they visit and follow them to flowers and then to the roost or 
perhaps directly to the roost site. They might also seek out pollen 


184 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


plants and follow experienced butterflies from there to the roost. 
However, it is not likely that a new individual would locate a plant 
like A6 easily without following other butterflies, since Anguria are 
generally inconspicuous, at least to human observers. Later foraging 
from the roost by experienced individuals may not involve the same 
following behavior which led to the roost’s discovery. Indeed, we did 
not observe following between established roost mates. 

These brief observations indicate the feasibility of relating roost 
membership to foraging behavior, interindividual interactions, and 
predation of individually numbered Heliconius butterflies in the field. 
We suspect that the major hypotheses for site-constant gregarious 
roosting (predator protection versus information center) in Heliconius 
will be difficult to clearly distinguish since, regardless of the reason 
for the evolution of the habit, other advantages to communal roosting 
may arise secondarily. The most certain conclusion is that Heliconius 
roosting behavior remains one of the major mysteries of lepidopteran 
biology and as such, deserves further study. 


ACKNOWLEDGMENTS 


We thank E. Gonzalez, D. Navarro, G. Perez, A. Estrada and the staff of the Estacion 
de Biologia Tropical “Los Tuxtlas” UNAM in Veracruz, Mexico for their assistance and 
hospitality during this study. D. Harvey helped with fieldwork. The manuscript has 
benefitted from comments by J. Mallett and P. DeVries. 


LITERATURE CITED 


BENSON, W. W. 1971. Evidence for the evolution of unpalatability through kin se- 
lection in the Heliconinae (Lepidoptera). Amer. Nat. 105:213-226. 

DUNLAP-PIANKA, H., C. L. BoGcs & L. E. GILBERT. 1977. Ovarian dynamics in 
Heliconiine butterflies: Programmed senescence versus eternal youth. Science 197: 
487-490. 

EHRLICH, P. R. & L. E. GILBERT. 1973. Population structure and dynamics of the 
tropical butterfly Heliconius ethilla. Biotropica 5:69-82. 

GaADGIL, M. 1972. The function of communal roosts: Relevance of mixed roosts. Ibis 
114:531—-533. 

GILBERT, L. E. 1975. Ecological consequences of a coevolved mutualism between 
butterflies and plants. In L. E. Gilbert and P. R. Raven (eds.). Coevolution of 
animals and plants. University of Texas Press, Austin. 

SINGER, M. C. & P. WEDLAKE. 1981. Capture does affect probability of recapture in 
a butterfly species. Ecological Entomology (in press). 

TURNER, J. R. G. 1975. Communal roosting in relation to warning coloration in two 
heliconiine butterflies (Nymphalidae). J. Lepid. Soc. 29:221-226. 

WARD, P. & A. ZAHAVI. 1973. The importance of certain assemblages of birds as 
“information-centres”’ for food-finding. Ibis 115:517-534. 

YOUNG, A. M. 1978. A communal roost of the butterfly Heliconius charitonius L. in 
Costa Rican premontane tropical wet forest (Lepidoptera: Nymphalidae). Entomol. 
News 89:235-243. 

& M. E. CAROLAN. 1976. Daily instability of communal roosting in the neo- 

tropical butterfly Heliconius charitonius (Lepidoptera: Nymphalidae). J. Kansas 

Entomol. Soc. 49:346-359. 


Journal of the Lepidopterists’ Society 
36(3), 1982, 185-191 


EVOLUTIONARY STUDIES ON CTENUCHID MOTHS OF THE 
GENUS AMATA: 2. TEMPORAL ISOLATION AND NATURAL 
HYBRIDIZATION IN SYMPATRIC POPULATIONS OF 
AMATA PHEGEA AND A. RAGAZZII 


V. SBORDONI,! L. BULLINI,”? P. BIANCO,’ R. CIANCHI,?” 
E. DE MATTHAEIS! AND S. FORESTIERO! 


ABSTRACT. Amata phegea and A. ragazzii are two sibling species which occur 
sympatrically in several areas of Central and Southern Italy. The occurrence of certain 
enzyme loci, electrophoretically diagnostic (PGM, HK, EST-5), allowed the correct 
identification of all individuals, including hybrids. Population studies in the field car- 
ried out in sympatric areas of Central Italy revealed the occurrence of seasonal dis- 
placement in the flight period acting as a premating isolating mechanism between the 
two species, in spite of a partial overlapping. Premating behavior of homospecific and 
heterospecific pairs investigated in the laboratory showed the existence of isolating 
mechanisms also at the ethological level, which lowers the mating success of the het- 
erospecific pairs. The frequency of hybrids in different localities varies from 0 to 0.053; 
the highest rates of hybridization were found in biotopes recently altered by man. 
Hybrid frequencies, similar in larval and adult stages, suggest a normal viability of the 


hybrids. 


The ctenuchid moths Amata phegea (Linnaeus, 1758) and A. ra- 
gazzii (Turati, 1917) are two sibling species, so far scarcely investi- 
gated from the genetic, ecological, ethological and zoogeographical 
points of view. A. phegea is widely distributed in Central Western 
Europe (with the exception of the Iberian peninsula) and is particu- 
larly abundant and widespread in Italy; while A. ragazzii is endemic 
in Central Souther Italy. The latter species has to date been ob- 
served in Calabria, Campania, Lucania, Molise, Lazio and Umbria. 

The two species presumably diverged through geographical isola- 
tion in warm refugia during the Pleistocene. They successively ex- 
panded their range in the post-glacial era following the spread of the 
deciduous broad-leaved forest. Areas of sympatric occurrence have 
been identified in the Alban Hills, in the Fioio Valley (Simbruini 
Mountains), near Leonessa (Rieti), at S. Polo dei Cavalieri (Sabine 
Mountains), at Mount Faito (Campania), at Viggiano (Lucanian Ap- 
pennines) and in other areas of Central and Southern Italy. 

At the morphological level identification of the adult of the two 
species may be difficult, particularly in females. On the other hand, 
allozyme separation by electrophoresis enables us to identify all in- 
dividuals, both larvae and adults (Bullini et al., 1981). Using this 


1 Institute of Zoology, University of Rome. 
2 Institute of Genetics, University of Rome. 
3 Institute of Zoology, University of L’Aquila. 


186 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


method we were able to discover some hybrids in the overlapping 
areas although no evidence of introgression has been detected. This 
seems to indicate the existence of highly effective isolating mecha- 
nisms. 

We are investigating these mechanisms both in the laboratory and 
in the field. In this paper we will present data showing patterns of 
temporal displacement in the flight period of the two species, prob- 
ably representing an effective premating isolating mechanism; a fur- 
ther premating mechanism is also reported operating at the etholog- 
ical level. In addition, we will present data on the occurrence of natural 
hybridization in some areas. 


STUDY SITES, MATERIALS AND METHODS 


As previously reported, a number of localities have now been iden- 
tified in Central Italy where A. phegea and A. ragazzii occur sym- 
patrically. In most of them the phenology of the two species was 
investigated by frequent observations over several years. More de- 
tailed information came from five localities near Rome, where peri- 
odic observations and counts were carried out. 

The first locality is situated at a height of 850 m near Camerata 
Nuova, a village 50 km north east of Rome at the foot of the Simbruini 
Mountains. This site is characterized by a Quercetum association and 
represents the middle-lower part of the biotope described in detail 
by Sbordoni et al. (1979). The remaining four sites, namely Zagarolo, 
San Cesareo, Rocca Priora and Monte Cavo, were selected along an 
altitudinal transect 14 km long from 300 to 900 m above sea level 
within an area including the Alban Hills about 30 km south east of 
Rome. Most of this area is characterized by chestnut woods. 

Observations were carried out from 1967 to 1980; reported data 
refer to 1974, which can be considered representative of the overall 
period. Direct counts of adult moths were employed to describe the 
phenology of A. phegea and A. ragazzii. Mark-release-recapture 
methods were also utilized to estimate absolute populations sizes of 
the two species (see Sbordoni et al., 1979). 

Starch gel electrophoresis applied to three diagnostic enzyme loci: 
phosphoglucomutase (Pgm), hexokinase (Hk) and esterase (Est-5) was 
utilized to identify morphologically doubtful specimens and to dis- 
cover hybrids (see Fig. 3). Electrophoretic techniques were, with mi- 
nor modifications, those described by Ayala et al. (1972) for phos- 
phoglucomutase and hexokinase, and by Selander et al. (1971) for 
esterase. 


VOLUME 36, NUMBER 3 : 187 


PERCENTAGE OF TOTAL FOR EACH SPECIES 


10 15 
JoeeUb oN) JE Ue a eeay, AS USER GRURS cali 


Fic. 1. Phenology of adult populations of Amata phegea (black circles) and A. 
ragazzti (white circles) at Camerata Nuova, Simbruini Mountains, 850 m, showing tem- 
poral displacement in the flight period. 


RESULTS AND DISCUSSION 
Temporal Displacement in Flight 


By scattared observations and preliminary collection data we noted 
that the two species emerge at different times. This trend is clearly 
apparent from the data reported here. 

At Camerata Nuova Amata phegea emerges about one month before 
A. ragazzii (Fig. 1). In the Alban Hills the emergence dates of the 
two species are separated by about 20 days, regardless of altitude (Fig. 
2). Similar phenological data were observed in other localities in Cen- 
tral Italy. The degree of overlapping depends on the relative popu- 
lation sizes of the two species, which vary according to the localities. 
At Camerata Nuova A. phegea greatly outnumbers A. ragazzii and the 
ratio between the population sizes is approximately 30:1 (Sbordoni et 
al., 1979). More balanced situations were found in the localities of 
the Alban Hills. Here, consistent overlapping is limited to 10-15 days. 

In both species males are first to emerge. Females are generally 
inseminated a few hours after their emergence (Stauder, 1927; Ob- 
raztsov, 1941). Female sex attractants are probably related to the anal 
papillae, which are rhythmically displayed by the virgin females (Ob- 
raztsov, 1966). Several males generally attempt to mate a single virgin 
female (Rasetti and Rasetti, 1921). Among adults the sex ratio is def- 


188 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


ZAGAROLO 300m S.CESAREO 450m 


5 10 15 20 25 5 10 15 20 «425 


5 10 15 20 25 6 10 15 20 25 
) UNE UIE Y. JUNE UP RY, 


ROCCA PRIORA 650m MONTE CAVO 900 m 


ok 


5 10 11% 20 25 5 10 15 20 25 5 10 18 20 25 5 10 15 20 25 
J UNE J Wok V JUNE Ho (WY tb MY 


PERCENTAGE OF TOTAL FOR EACH SPECIES 


Fic. 2. Phenologies of adult populations of Amata phegea (black circles) and A. 
ragazzii (white circles) in 4 localities along an altitudinal transect south east Rome. 


initely in favor of the males (80-85% in both species). According to 
these behavioral traits and the sex ratio, it seems improbable that a 
female of A. phegea could be inseminated by a male of A. ragazzii. 
On the other hand, old males of both species show an erratic behavior, 
and they tend to wander from their usual habitat in forest edges, open 
woods, etc. to open fields; also, this behavior limits the opportunities 
of mating between males of A. phegea and emerging females of A. 
TAZAZZU. 

These observations led us to conclude that seasonal displacement 
may act as a premating isolating mechanism between A. phegea and 
A. ragazzii, although partial temporal overlapping occurs. This barrier 
would be particularly effective if monogamy were present in both 
species. However, this latter point needs to be tested. 

Similar temporal displacement occurs also in other sympatric com- 
binations of related Amata species. We observed this phenomenon 
in Istria and Dalmatia between A. marjana (Stauder, 1913) and A. 
phegea. In such instances, A. marjana is the first to fly, and the emer- 
gences of the two species are separated by about one month. 


Premating Behavior 


Besides the temporal displacement, an ethological isolating mech- 
anism was also detected by preliminary laboratory experiments. A. 


VOLUME 36, NUMBER 3 189 


2 3 4 56°78 9 0 1 2 BK 


Fic. 3. Phosphoglucomutase (PGM) electrophoretic patterns of Amata phegea, A. 
ragazzii and their hybrid. A. ragazzii (specimens 14 and 10-14, from left to right) 
shows an anodic band migrating faster than A. phegea (specimens 5 and 7-9), whereas 
the hybrid (specimen 6) shows the two bands characteristic of the parental species. 


phegea males isolated with conspecific virgin females are strongly 
attracted and approach the partner with a characteristic zig-zag flight. 
When the male touches the female she stops flying and the male lands 
on her. He repeatedly touches the costal area of her anterior wings 
with his antennae; the female, keeping her wings spread, raises her 
abdomen. Then copulation takes place. When A. ragazzii males are 
isolated with virgin A. phegea females, precopulatory behavior always 
begins later. Furthermore, the female often doesn’t stop when touched 
and moves away. This generally interrupts a male's precopulatory 
behavior. Even when copulation takes place, the female frequently 
lays unfertilized eggs. 


Natural Hybridization Between A. phegea and A. ragazzii 


In this section we will present data demonstrating the occurrence 
of natural hybrids between the two Amata species occurring sym- 
patrically in areas of Central Italy. 

As reported in the materials and methods section, detection of hy- 
brids by electrophoresis was made at some enzyme loci, which are 
diagnostic for A. phegea and A. ragazzii (Pgm, Hk, Est-5). At these 
loci no common allele is shared between the two Amata species stud- 
ied. Hybrids are characterized by heterozygous patterns at each of 
these loci. 

In Fig. 3 an example of a zymogram is shown, illustrating the pat- 
tern of A. phegea, A. ragazzii and their hybrid, at the Pgm locus. 

Table 1 shows the numbers of A. phegea, A. ragazzii and hybrids 
in samples from some sympatric areas in Central Italy. Samples tab- 
ulated were collected during periods of temporal overlapping of the 
two species. However, if samples are collected over the whole period 
of flight of the two species, rates of hybridization may appear lower, 


190 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 1. Rates of natural hybridization between A. phegea and A. ragazzii, as 
revealed by electrophoresis on samples of adult moths from some sympatric areas of 
Central Italy. 


Number of specimens scored Relative 
 ——___ frequency 
Study site Date of sampling A. phegea A. ragazzii Hybrids of hybrids 
Camerata Nuova 3-23 July 1974 ive 21 1 0.005 
Camerata Nuova 7 July—6 August 1975 891 99 3 0.003 
Leonessa 25 July-8 August 1976 205 25 1 0.004 
Montoro 25 June-23 July 1979 53 50 1 0.009 
San Polo 4-19 July 1976 71 114 1 0.005 
San Cesareo 8 July 1974 q 12 il 0.053 
Tuscolo 8 July 1974 1 11 = = 
Monte Compatri 16 June-9 July 1974 39 10 — = 
Monte Porzio Catone 3-30 July 1975 10 202 il 0.004 
Monte Cavo 30 July 1975 40 58 2 0.02 


because all the hybrids detected were in flight relatively later, to- 
gether with A. ragazzii. 

Table 2 shows the data obtained from samples of larvae collected 
in three sites of the Alban Hills. 

A comparison between the two tables do not reveal substantial dif- 
ferences in the frequency of hybrids between adult and larval stages 
from the same locality, suggesting normal viability of hybrids. 

The frequency of hybrids varies from 0 to 0.053, but several values 
are around 0.004. The high rate of hybridization detected at San Ce- 
sareo, both from larval and adult samples observed in two distinct 
generations, may be tentatively related to the man-made alteration of 
their biotope. At the collection sites a wide zone of chestnut wood 
was replaced by Rubus sp. 

Cases of sympatric hybridization are often associated with habitat 
alteration (Woodroof, 1973). In the case of A. phegea and A. ragazzii 
habitat alteration could affect the rate of hybridization and even the 
occurrence of sympatry between the two species. This working hy- 
pothesis requires further investigation. 


TABLE 2. Rates of natural hybridization between A. phegea and A. ragazzii, as 
revealed by electrophoresis on samples of larvae from some sympatric areas of Central 
Italy. 


Number of specimens scored Relative 


SS  ieaieney 

Study site Date of sampling A. phegea A. ragazzii Hybrids of hybrids 
San Cesareo February 1975 50 107 6 0.038 
Tuscolo 27 January 1975 5 113 II 0.008 


Monte Compatri February 1975 62 65 — — 


VOLUME 36, NUMBER 3 191 


LITERATURE CITED 


AYALA, F. J., J. R. POWELL, M. L. TRACEy, C. A. MOURAO & S. PEREZ-SALAS. 1972. 
Enzyme variability in the Drosophila willistoni group. IV. Genic variation in nat- 
ural populations of Drosophila willistoni. Genetics 70:113-139. 

BULLINI, L., R. CIANCHI, C. STEFANI & V. SBORDONI. 1981. Biochemical taxonomy 
of the Italian species of the Amata phegea complex (Ctenuchidae, Syntominae). 
Nota Lepid. 4: in press. 

OBRAZTSOV, N.S. 1941. Die Artdifferenzierung der phegea-Gruppe des Subgenus 
Syntomis O. der europaischen Fauna. Univ. Kijev. Acta Mus. Zool. 1:103-164. 
1966. Die palaearktischen Amata-Arten (Lepidoptera, Ctenuchidae). Veroff. 

Zool. Staatssamml. 10:1-383. 

RASETTI, E. & F. RASETTI. 1921. Note entomologiche. Boll. Soc. Entomol. Ital. 
03: 19-27. 

SBORDONI, V., L. BULLINI, G. SCARPELLI, S. FORESTIERO & M. RAMPINI. 1979. Mim- 
icry in the burnet moth Zygaena ephialtes: population studies and evidence of a 
Mullerian-Batesian situation. Ecol. Entomol. 4:83-93. 

SELANDER, R. K., M. H. SMITH, S. Y. YANG, W. E. JOHNSON & J. B. GENTRY. 1971. 
Biochemical polymorphism in the genus Peromyscus. I. Variation in the old field 
mouse (Peromyscus polionotus). Studies in Genetics, 6, Univ. Texas Publ. 
7103:49-90. f 

STAUDER, H. 1927. Uber Zucht suditalienischer Syntomiden. Lepid. Rundsch. 1:57-59. 

WoopROOF, D.S. 1973. Natural hybridization and hybrid zone. Syst. Zool. 22:213-217. 


Journal of the Lepidopterists’ Society 
36(3), 1982, 192-199 


COCOONS OF CALLOSAMIA PROMETHEA (SATURNIIDAE): 
ADAPTIVE SIGNIFICANCE OF DIFFERENCES IN 
MODE OF ATTACHMENT TO THE HOST TREE 


G. P. WALDBAUER AND J. G. STERNBURG 


Department of Entomology, University of Illinois, 
320 Morrill Hall, 505 S. Goodwin, Urbana, Illinois 61801 


ABSTRACT. The cocoons of Callosamia promethea (Drury) (Saturniidae) are 
wrapped in a leaf and hang from a twig of the host tree by a silken peduncle that 
sheathes the leaf petiole and by a silken anchor that sheathes a variable length of the 
woody twig. It is proposed that in winter the cocoon’s height above ground tends to 
protect it from mice and that its flexible attachment to a thin twig tends to protect it 
from woodpeckers. The anchor is usually about 2 cm long, but on thin twigs it may be 
much longer, sometimes extending past the next fork of the branch. The extension of 
the anchor seems superfluous on most of promethea’s hosts, trees with simple leaves 
where anchoring the petiole to the adjoining twig is sufficient to assure the cocoon’s 
continued attachment to the tree after leaf fall. However, some of promethea’s hosts, 
the ashes (Fraxinus spp.), have compound leaves, and on these trees the cocoon will 
fall with the leaves in autumn unless the anchor is extended from the leaflet petiole 
up the rachis to encircle the adjoining woody twig. 


Pupae of Callosamia promethea (Drury) (Saturniidae) overwinter 
in cocoons that dangle freely from a strong flexible silken peduncle 
anchored to a twig of the host tree (Fig. 1). In spinning the cocoon 
the larva first rolls a leaf along its midrib, fastens it at the margins, 
and lines it with silk to form an open-ended tube. It then spins a 
peduncle and anchor that are continuous with the lining of the leaf 
tube, the peduncle sheathing the leaf petiole and the anchor sheath- 
ing a variable length of the adjoining twig. Finally, the larva reenters 
the leaf-tube to spin a tough double-walled cocoon with a valve for 
the emergence of the adult at its top, where the petiole joins the leaf 
blade (Haskins & Haskins, 1958). The cocoons are usually fixed to 
thin terminal twigs well above the ground at the periphery of the 
tree’s crown. They do not fall with the leaves in autumn. The envel- 
oping leaf usually weathers away in winter, but the peduncle and 
anchor remain intact, securely attaching the cocoon to the tree (Fer- 
guson, 1972). 

The cocoon’s height above ground probably tends to protect it from 
mice and its flexible attachment to a thin terminal twig, from wood- 
peckers. Accordingly, we present data on the predation pressure on 
promethea moth pupae, comparing it with the predation pressure, 
determined in other studies, on the pupae of a sympatric saturniid, 
Hyalophora cecropia (L.), whose larger cocoon often occurs in the 
same habitat but is immovably fixed to the stem or branch of a woody 
plant, usually near ground level (Scarbrough et al., 1972a). We also 


VOLUME 36, NUMBER 3 193 


Fic. 1. Callosamia promethea cocoons showing the variations in the mode of at- 
tachment to the host tree. A, a short anchor, the usual mode of attachment; B, an 
extended anchor that does not reach past a fork; C, an anchor that extends past the next 


fork of the branch. 


present data on the extent of the promethea cocoon’s anchor. It is 
usually short but may be long, sometimes even extending up the twig 
past the first fork to sheathe a more proximal and thicker part of the 
branch (Fig. 1, Table 2). 


MATERIALS AND METHODS 


We collected promethea cocoons in east central Illinois from Dan- 
ville south to Interstate Highway 70, and in northwestern Indiana 
from I-70 north to Medaryville. All were found on black cherry (Pru- 
nus serotina Ehrh.) or sassafras (Sassafras albidum Nees.) saplings 
that were seldom more than 3 to 4 m tall and were usually in fence 


rows in agricultural areas. 


194 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


TABLE 1. Total Callosamia promethea cocoons collected, and the number and per- 
cent dead from various causes including unidentified parasites. Cocoons from Indiana 
were collected on 4 March 1972, and cocoons from Illinois were collected on 28 March 
1972. 


Cause of death 


Total Unknown Parasite Woodpecker Mouse 

lected’ Nor) % No.) Sagas JIN Oncaea 
Medaryville, IN 2000 ies: 3 4 2.0 9), «4:5 sons 
Reynolds, IN 121 6) a0 0) 0 OF0 La OS 
Charleston, IL 91 5 B0r0n) 200 LL 22R0) AN 4740 Oe a0) 
Total or percent of total AND T2388 6324 5.8. 13.2 SQ 322205 


Random samples of promethea cocoons for estimates of predation 
pressure were collected at the localities and on the days in March 
shown in Table 1. Most predation had probably occurred prior to 
collection; Sternburg et al. (1981) found that 82.4% of the woodpecker 
attacks on cecropia moth cocoons had occurred by 4 March. Non- 
random samples of cocoons for determining the dimensions of the 
anchors and of the supporting twigs were clipped from trees with the 
anchor intact on 29 December 1969 near Medaryville, Indiana. We 
tried to find as many as possible of the relatively scarce cocoons with 
long anchors. 

Length was measured with a rule and diameter with a micrometer. 
Dimensions of the distal part of the anchor (Table 2) were analyzed 
with a one-way ANOVA followed by the Student-Newman-Keuls test 
(Sokal & Rohlf, 1969). The mean lengths of the distal portions of 
anchors extending past the first fork (Fig. 1) cannot be legitimately 
compared with each other or with shorter anchors because, by defi- 
nition, the lengths of the former are determined by the distance to 
the fork, while the lengths of the latter are not so determined. 


RESULTS AND DISCUSSION 


Thirteen (3.2%) of the promethea pupae had been killed by wood- 
peckers and only two (0.5%) by mice (Table 1). We found predation 
to be similarly light on several thousand cocoons collected in ten 
years at or near the same localities. However, on 1 March 1982, after 
two months of unusually deep snow cover, about 48% of the prome- 
thea cocoons that we found along 8 km of roadside near Medaryville 
had been attacked by woodpeckers, although promethea cocoons in 
nearby areas had not been attacked. 

The similarity of the damage on these promethea cocoons to dam- 
age of known origin on cecropia cocoons leaves no doubt that the 


195 


VOLUME 36, NUMBER 3 


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196 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


former were attacked by woodpeckers and mice. The downy (Den- 
drocopus pubescens (L.)) and the hairy (D. villosus (L.)) woodpeckers 
pierce the cocoon, making a small hole through which their barbed 
tongues remove the viscous pupal contents (Waldbauer et al., 1970). 
Both of them are common in promethea’s habitat in winter (Bent, 
1964). The mice Peromyscus maniculatus (Wagner) and P. leucopus 
(Raf.) remove the entire pupa through a large hole which they chew 
in the cocoon (Scarbrough, 1970; Scarbrough et al., 1972b). They are 
also common in promethea’s habitat (Hoffmeister & Mohr, 1972). We 
found only one type of damage that may have been caused by another 
predator. A few cocoons were crimped and the pupae partly crushed, 
as if pinched by the bill of a bird. One of us (J.G.S.) saw a blue jay 
(Cyanocitta cristata L.) in January pinch and then desert a promethea 
cocoon that was later found to contain only exuviae. 

Although cecropia is largely urban (Scarbrough, 1970) and prome- 
thea is largely rural (Sternburg & Waldbauer, unpublished), both 
species feed on black cherry and may occur in the same rural fence 
rows. We have found that cecropia cocoons often fall prey to wood- 
peckers and mice in this habitat. 

Although we do not have comparable predation data for these two 
species from the same area, there is no doubt that both woodpeckers 
and mice generally take a far heavier toll of cecropia than of prome- 
thea. In both urban and rural areas woodpeckers regularly kill about 
90% of the cecropia pupae in cocoons 45 cm or more above the ground 
(Waldbauer & Sternburg, 1967a, b). In rural areas mice destroy as 
many as 60% of the cecropia pupae near ground level (Scarbrough, 
1970; Scarbrough et al., 1972b). 

The far lower level of predation on promethea cocoons suggests 
that their greater height above the ground, flexible attachment to a 
thin twig, and perhaps their smaller size may be adaptive responses 
to predation by vertebrates. Although Peromyscus leucopus are some- 
what arboreal, they rarely attack high cecropia cocoons (Scarbrough 
et al., 1972b) or promethea cocoons (Table 1). Woodpeckers may perch 
directly on the large immovable cecropia cocoons (Waldbauer et al., 
1970), but they are probably reluctant to perch on the far smaller and 
free swinging promethea cocoon. Nielsen (1977) saw a downy wood- 
pecker hang from a promethea cocoon as it pierced the pupa, but our 
data (Table 1) indicate that this is uncommon. The larger hairy wood- 
pecker may find it even more difficult to perch on promethea cocoons 
than does the smaller downy. The thin twigs that support promethea 
cocoons may not be secure perches for woodpeckers. Even if a wood- 
pecker does find a perch near a cocoon, it may not be able to pierce 
it because the cocoon, hanging by its flexible peduncle, swings away 


VOLUME 36, NUMBER 3 197 


when it is pecked. About 77% of the woodpecker-attacked cocoons 
listed in Table 1 had been pierced down through the valve into the 
head of the pupa. About 34% of a sample of 38 woodpecker-attacked 
promethea cocoons collected in 1982 had been similarly attacked. 
Woodpeckers may tend to attack in this way because the force of a 
peck directed down into the valve does not cause the cocoon to swing 
away. 

The length of the anchor of promethea cocoons varies greatly (Table 
2). Those with short anchors (2 cm or less) are most common; those 
with long anchors (up to 19 cm) that do not extend proximad past the 
first fork in the twig are much less common; and those with long 
anchors (up to 15 cm total length) that do extend past the first fork are 
the least common. Note that the numbers in Table 2, not based on 
random samples, do not reflect the relative abundance in nature of 
these three anchor types. 

The data in Table 2 indicate that a cue associated with the diameter 
of the supporting twig stimulates promethea larvae to spin an extend- 
ed anchor. Cocoons with anchors extending past the first fork of the 
twig were on the thinnest twigs, those with long anchors not extend- 
ing past the fork were on somewhat thicker twigs, and those with 
short anchors (2 cm or less) were on the thickest twigs. The mean 
diameters of twigs in each category are significantly different on both 
black cherry and sassafras (Table 2). Whether spinning larvae extend 
the anchor in response to the relative thinness of the twig, or to some 
other property associated with thinness, perhaps greenness, cannot 
be determined from the data at hand. While green twigs are probably 
thinner than woody twigs on the same tree, they are also softer; and 
promethea larvae may extend the anchor in response to a relatively 
soft-textured supporting twig. 

It is reasonable to hypothesize that the extension of the anchor is 
intended to prevent the cocoon from falling to the ground where it 
can be found by mice. However, extension of the anchor on black 
cherry or sassafras, trees with simple leaves, seems superfluous since 
even a short attachment to the adjoining woody twig is sufficient to 
prevent the cocoon from falling with the leaves in autumn. Woody 
twigs seldom fall spontaneously from these trees, and there appears 
to be no present danger from predators that might be better able to 
sever a thin twig than a thick one. 

We suggest that the extension of the anchor is actually an inappro- 
priate manifestation of a behavior that evolved as an accommodation 
to host plants with compound leaves. The rachis of a compound leaf 
is green and softer than a woody twig, and it is shed in winter. The 
anchor must extend up the rachis to the woody twig to keep the co- 


198 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


coon from falling with the leaves. The promethea host plants listed 
by Ferguson (1972) and Tietz (1972) include only one group with 
compound leaves, the ashes (Fraxinus spp.). We do not find prome- 
thea cocoons on ash, but Comstock and Comstock (1916) found them 
to be abundant on ash. Promethea may have been more common on 
ash, or may even have used other plant species with compound leaves, 
when the capacity to extend the anchor evolved. This behavior might 
also be adaptive on any of the willows (Salix spp.) that shed some of 
their woody twigs. Ferguson (1972) and Tietz (1972) list willows as 
promethea food plants, and Vestal (1913) found their cocoons abun- 
dantly on willow near Havana, Illinois. 

Cocoons on wild cherry are on thinner twigs, regardless of the length 
of the anchor, than are cocoons on sassafras (Table 2); these differ- 
ences in mean diameter are significant (P < 0.05, Student's t test). If 
we are correct in our conjecture that promethea larvae respond to 
green twigs by spinning a long anchor, then these differences reflect 
only the greater mean diameter of sassafras twigs. If, on the other 
hand, the larvae actually respond to thinness per se, then these dif- 
ferences suggest that the larvae compare the thickness of the sup- 
porting twig with the thinner leaf petiole or with thicker twigs tra- 
versed enroute to the spinning site. 

Pammer (1966) found that another saturniid, Samia cynthia Drury, 
is adapted to cope with compound leaves. Cynthia larvae feed on 
Ailanthus altissima (Mill.) Swingle (Ferguson, 1972), a tree with large, 
singly compound, deciduous leaves. The caterpillars spin cocoons 
that, like promethea cocoons, are wrapped in a leaflet and have a 
peduncle extending up the petiole. Larvae of the summer generation 
anchor the leaflet only to the rachis; they emerge as adults before the 
leaves fall. Larvae of the overwintering generation, however, ensure 
their continued attachment to the tree by extending the anchor up 
the rachis to the adjoining woody twig. 

Promethea is partly bivoltine in the southern part of our collecting 
area, but it is yet to be determined if second generation larvae are 
more likely to extend the anchor than are first generation larvae. 


LITERATURE CITED 


BENT, A. C. 1964. Life histories of North American woodpeckers. Dover Publications, 
New York. 334 pp. 

COMSTOCK, J. H. & A. B. Comstock. 1916. A manual of the study of insects. 14th 
ed. Comstock Pub. Co., Ithaca, N.Y. 701 pp. 

FERGUSON, D.C. 1972. Bombycoidea (in part). Pp. 246-259 in Dominick et al. (eds.). 
The moths of America north of Mexico. Fascicle 20.2B. Classey, London. 

HASKINS, C. P. & E. F. HASKINS. 1958. Note on the inheritance of behavior patterns 
for food selection and cocoon spinning in F, hybrids of Callosamia promethea x 
C. angulifera. Behaviour 13:89-95. 


VOLUME 36, NUMBER 3 199 


HOFFMEISTER, D. F. & C. O. MouR. 1972. Fieldbook of Illinois mammals. Dover 
Publications, New York. 233 pp. 

NIELSEN, M. C. 1977. Woodpecker feeding on Callosamia promethea (Saturniidae) 
cocoon. J. Lepid. Soc. 31:148-149. 

PAMMER, E. 1966. Auslosung und Steuerung des Spinnverhaltens und der Diapause 
bei: Philosamia cynthia Dru. (Saturniidae, Lep.). Z. vergl. Physiol. 53:99-113. 
SCARBROUGH, A. G. 1970. The occurrence of Hyalophora cecropia (L.) as related to 

urbanization. Ph.D. thesis, University of Illinois, Urbana. 217 pp. 

, ]. G. STERNBURG & G. P. WALDBAUER. 1972a. Selection of the cocoon spin- 

ning site by the larvae of Hyalophora cecropia (Saturniidae). J. Lepid. Soc. 31: 

153-166. 

, G. P. WALDBAUER & J. G. STERNBURG. 1972b. Response to cecropia cocoons 
of Mus musculus and two species of Peromyscus. Oecologia 10:137-144. 

SOKAL, R. R. & F. J. ROHLF. 1969. Biometry. W. H. Freeman & Co., San Francisco. 
776 pp. 

STERNBURG, J. G., G. P. WALDBAUER & A. G. SCARBROUGH. 1982 (1981). Distribution 
of cecropia (Saturniidae) in central Illinois: A study in urban ecology. J. Lepid. 
Soc. 35(4):304-320. 

TiETZ, H. M. 1972. An index to the life histories, early stages and hosts of the Mac- 
rolepidoptera of the continental United States. Vol. I1:537-1041. 

VESTAL, A. G. 1913. An associational study of Illinois sand prairie. Bull. Illinois State 
Lab. Nat. Hist. 10:1-96. 

WALDBAUER, G. P. & J.G. STERNBURG. 1967a. Host plants and the locations of baggy 
and compact cocoons of Hyalophora cecropia (Lepidoptera: Saturniidae). Ann. 
Entomol. Soc. Amer. 60:97-101. 

& 1967b. Differential predation on cocoons of Hyalophora cecropia 

(Lepidoptera: Saturniidae) spun on shrubs and trees. Ecology 48:312-315. 

: , W. G. GEORGE & A. G. SCARBROUGH. 1970. Hairy and downy wood- 
pecker attacks on cocoons of urban Hyalophora cecropia and other saturniids (Lep- 

idoptera). Ann. Entomol. Soc. Amer. 63: 1366-1369. 


Journal of the Lepidopterists’ Society 
36(3), 1982, 200-206 


NOTES ON THE ACOUSTIC SIGNALS OF A 
NEOTROPICAL SATYRID BUTTERFLY 


STEPHANIE KANE 


Department of Zoology, The University of Texas, 
Austin, Texas 78712* 


ABSTRACT. Acoustical signalling is documented for the satyrid, Pharneuptychia 
nr. pharnabazos. The clicking sounds are produced during specific flight behaviors. 
The sounds have been identified from males only to date, but no structures or mech- 
anisms for producing the sounds were identified. 


Sound production in butterflies is a sporadic phenomenon, largely 
reported in the genus Hamadryas (Nymphalidae) (Darwin, 1871; 
Fruhstorfer, 1924; Ehrlich & Ehrlich, 1961; Ross, 1963). Audible 
clicking sounds in flight are characteristic of this group, hence the 
common name of “crackers.” The nature of their sound producing 
mechanism is as yet unresolved (Swihart, 1967). Acoustic signalling 
has also been observed in Neptis hylas (Linnaeus) (Nymphalidae) 
(Scott, 1968), which produces a slow clicking sound by snapping to- 
gether its forelegs while in a resting position. This paper presents the 
first documented evidence of acoustic signalling in a satyrid butterfly. 
The signals consist of bursts of clicking sounds emitted during ritu- 
alized flight behavior. The butterfly was identified as Pharneuptychia 
nr. pharnabazos by Dr. Keith Brown; however, the specimens appear 
quite similar to the plates in Seitz, Volume 5 (Plate 48d) called Eup- 
tychia pharella (Butler) (Satyridae). For the purposes of discussion, I 
will call it Pharneuptychia nr. pharnabazos, although the identity is 
uncertain at this point. 


Field Site and Method 


Sound production was documented in the adult Pharneuptychia nr. 
pharnabazos between 0800 and 1000 h from 31 March to 4 April 1980 
near the town of Senador Pompeu in the state of Ceara in northeastern 
Brazil (6°S latitude, 39°W longitude). A recording of the acoustic sig- 
nals was made from 0900 h on 31 March with a Uher 4000L tape 
recorder, a Sony C-22 FET condenser microphone and parabola from 
a distance of 0.5 to 1.0 meter. Sound spectrograms were made on a 
Kay Electric Co., Type 6061-A sonagraph using FL-1, linear scale, 
and wide-band filter settings. A copy of the recording was deposited 
in the Cornell Library of Natural Sounds and the Arquivo Sonoro 
Neotropical (cut S. Kane no. 17/1) of the Universidade Esta- 
dual de Campinas, Sao Paulo, Brasil. Voucher specimens were de- 
posited in the American Museum of Natural History. 


* Now in Dept. of Anthropology (same university). 


201 


VOLUME 36, NUMBER 3 


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202 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Table |. Bioacoustic Data 
signal signal spectro- frequency range click rate of 
# duration gram of spectrogram spectrogram 
(sec) | (kHz) (click/sec.) 

| | A 4-13 I9.6 

2 4 B Sez 5 28.4 

€ 6 715 I2.8 

3 2 D ZS 17.2 


Fresh fruit of the Juazeiro tree (Zizyphus juazeiro Mart., Rhamna- 
ceae) was cut up and distributed on rocks about 30 cm apart on the 
ground in a sunlit forest clearing. This attracted P. nr. pharnabazos 
and two other common fruit visiting species, Hamadryas ferentina 
Godart and Anaea moretta Druce (Nymphalidae). Intra- and inter- 
specific behavioral interactions were thereby observed with facility. 
Similar behavioral interactions were also observed along paths, other 
open woodland areas, and near fruiting trees. , 


Bioacoustic Analysis 


Original recordings were made at 7% ips. The spectrograms rep- 
resent 1.2 s segments of recorded signals in the range of 800 to 16,000 
Hz (see Fig. 1). The total recording duration is 32 s and consists of 
three signals from one or two individuals. The mean signal duration 
(n = 3) = 2.33 = 1.53" s. Mhe mean inter-signall durations —s) a 
12.5 + Os. A signal is defined as one or more groups of clicks sepa- 
rated by neighboring groups by 12.5 s. The click is a broad band sound 
pulse with a range of frequencies between 4.25 kHz to 14.5 kHz for 
the greatest part of the amplitude of sound. Mean click rate per spec- 
trogram = 19.5 clicks/s (see Table 1). 

The sound producing mechanism is uncertain. It may be associated 
with wing beat, but this cannot be verified from the data presented, 
especially since, in some cases, two individuals may have been re- 
corded simultaneously. A file system could produce the sounds as in 
crickets (Alexander, 1967). Unfortunately, the specimens had been 
crushed by the time they were examined microscopically and neither 
sound-producing, nor sound-receiving organs were apparent. 


VOLUME 36, NUMBER 3 203 


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Fic. 2. Model of flight patterns of A) circling behavior and B) follow and chase 
behavior of 2 individual P. nr. pharnabazos butterflies. Relative strength of line indi- 
cates corresponding positions of individuals. Sounds are produced discontinuously 
throughout the movements. 


Behavioral Notes 


Auditory signals accompany ritualized flight behavior of two or three 
individuals. Sounds are produced discontinuously during circling and/ 
or during follow and chase behavior (see Fig. 3). P. nr. pharnabazos 
usually flies about 10 cm off the ground, going up to 20 and 30 cm 
during the circle dance. Of six individuals collected, all are males. 
Three of these were clicking at the time of capture. It is not known 
if females produce sounds. Intraspecific encounters of another satyrid, 
Erebia epipsodea Butler (Brussard & Ehrlich, 1970), include ascend- 
ing spiral flights and chasing behavior. These behaviors are also part 
of the intraspecific male territorial defense repertoires of Archonias 
tereas (Godart) (Pieridae) (Gilbert, 1968), Papilio zelicaon Lucas 
(Papilionidae) (Gilbert, pers. comm.; also described in Maynard Smith 
& Parker, 1976), and Pararge aegeria (Linnaeus) (Nymphalida) (Da- 
vies, 1978). Other genera of butterflies having male-male spiral chases 
are Adelpha (Nymphalidae), Epiphile (Nymphalidae), Catasticta 
(Pieridae), and some Cissia species (Satyridae) (pers. comm., Philip 
DeVries). 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Fic. 3. Upper and under sides of the voucher specimen Pharneuptychia nr. phar- 
nabazos. 


VOLUME 36, NUMBER 3 205 


The intra- and interspecific interactions described below strongly 
suggest that the acoustic signals of P. nr. pharnabazos accompany 
agonistic behavior in the presence of food and/or mates. These signals 
may also function as part of an intraspecific recognition code, main- 
taining contact between male and female individuals of a mobile group. 


Intraspecific encounters of Pharneuptychia nr. pharnabazos: 

1. Individual (A) approaches posed individual (B). A clicks briefly. 
A and B fly off, clicking while following and circling each other. Both 
pose again. 

2. Two males in flight. There is clicking during alternating circle 
and follow behavior. 

3. Two individuals (A, B) are posed. A third individual (C) ap- 
proaches A, clicking and circling, chases off C. 

4. Two individuals (A, B) encounter one another in flight. They fly 
soundlessly in the same direction until a third individual (C) appears. 
A separates from B and chases C, clicking. B and C pose near one 
another, flashing wings. 

5. Individual (A) is feeding. Individual (B) approaches and waits 1 
to 2 minutes. B then clicks and flies closely around A, crashing into 
A. B leaves and poses nearby for 2 minutes. B comes back and A is 
still feeding. B flies off to another piece of fruit some distance away 
but does not feed. 

6. Individual (A) is feeding. Individual (B) is posed close to A. B 
clicks as a third individual (C) approaches. C flies away. 


Interspecific interaction between Pharneuptychia nr. pharnabazos 
and Hamadryas ferentina: 

1. H. ferentina (H) approaches two individual P. nr. pharnabazos 
butterflies (A, B). A takes off and flies, clicking around H. A third 
individual P. nr. pharnabazos (C) joins A, both click and fly after H. 
B remains feeding. 


RESUMO 


Este trabalho apresenta a primeira evidéncia de sinalizacao acustica 
de uma borboleta da familia Satyridae, identificada como sendo Phar- 
neuptychia aff. pharnabazos do nordeste do Brasil. Os sinais consti- 
tuem-se em estalidos em séries, emitidos em ritual de voo entre dois 
ou trés individuos. Andlise bioactistica dos sons gravados e as descri- 
coes de comportamento estao incluidas. 


ACKNOWLEDGMENTS 


I would like to acknowledge the welcome and orientation I received from the Zo- 
ology Department of the Universidade Estadual de Campinas, Sao Paulo, Brazil, and 


206 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


the hospitality of the Sa Magalhaes family. I am grateful to Cornell Natural Sound 
Library for their loan of recording equipment. I appreciate the editorial criticism of 
earlier drafts of this manuscript by Lawrence Gilbert and Philip DeVries, and the photo 
of P. nr. pharnabazos also by Philip. I thank Lincoln Stoller for his butterfly flight path 
drawing. 


LITERATURE CITED 


ALEXANDER, R. D. 1967. Acoustical communication in arthropods. Ann. Rev. Ento- 
mol. 12:495-525. 

BRUSSARD, P. F. & P. R. EHRLICH. 1970. Adult behavior and population structure in 
Erebia episodea (Lepidoptera: Satyrinae). Ecology 51(5):880—-885. 

DARWIN, C. R. 1871. Descent of man. Modern Library, New York. 

Davies, N. B. 1978. Territorial defense in the speckled wood butterfly (Pararge 
aegeria): The resident always wins. Anim. Behav. 26: 138-147. 

EHRLICH, P. R. & A. H. EHRLICH. 1961. How to know butterflies. W. C. Brown, 
Dubuque, Iowa. 262 pp. 

FRUHSTORFER, 1924. In A. Seitz. The Macrolepidoptera of the World. Alfred Kernen, 
Stuttgart. 

GILBERT, L. 1968. Studies on butterfly populations in several wet forest localities of 
Costa Rica. Part II: Territorial behavior by male Archonias butterflies (Pieridae). 
Individual Research Project, Advanced Population Biology Course (Winter, 1968). 
Organization for Tropical Studies (unpublished). 

MAYNARD SMITH, J. & G. A. PARKER. 1976. The logic of assymetric contests. Anim. 
Behav. 24:159-175. 

Ross, G. N. 1963. Evidence for lack of territoriality in two species of Hamadryas 
(Nymphalidae). J. Res. Lepid. 2(4):241-246. 

ScoTT, F. W. 1968. Sounds produced by Neptis hylas (Nymphalidae). J. Lepid. Soc. 
22(4):254. 

SWIHART, S. L. 1967. Hearing in butterflies (Nymphalidae: Heliconius, Ageronia). J. 
Insect Phys. 13:469-476. 


Journal of the Lepidopterists’ Society 
36(3), 1982, 207-216 


RELATIONSHIPS BETWEEN PUPAL SIZE AND SEX IN 
GIANT SILKWORM MOTHS (SATURNIIDAE)! 


THOMAS A. MILLER, JERRY W. HIGHFILL,? AND WILLIAM J. COOPER?® 
U.S. Army Medical Bioengineering R&D Laboratory, Fort Detrick, Maryland 21701 


ABSTRACT. Weights and dimensions are given for pupae of Callosamia prome- 
thea (Drury), Eupackardia calleta (Westwood) and Hyalophora cecropia (Linnaeus). 
Significant sex-related differences were observed in sample means for all characteris- 
tics studied, except antenna length in C. promethea. Discriminant function equations 
were derived for predicting sex in individual pupae of E. calleta on the basis of weight 
and antenna width data; and individual pupae of H. cecropia on the basis of circum- 
ference and antenna width data. Reliable discriminant function equations could not be 
derived for determining sex in individual C. promethea pupae. Within species, sig- 
nificant differences were observed for male and female antenna surface areas. Between 
species, antenna length to width ratios did not differ significantly for individuals of the 
same sex. 


The ability to determine sex in lepidopterous pupae precludes the 
need to await adult emergence to identify individuals for breeding or 
experimentation, or to determine sex ratios or individual sexes. For 
lepidopterous pupae the dimensions of the antennae and the mor- 
phology of the genital openings have been the most reliable and widely 
used characteristics for determining sex. Other characteristics such as 
coloration, body size, and even behavior have been used for certain 
species. The fact that female pupae are generally larger than males 
has been noted by many lepidopterists, but such differences have not 
been quantified. (Mosher, 1914, 1916a, 1916b; Butt & Cantu, 1962; 
Solomon, 1962; Ehrlick et al., 1969; Villiard, 1969; Kean & Platt, 1973; 
Jennings, 1974; Mugegli, 1974). 

Mosher (1914, 1916a, 1916b) is the best available reference on sex- 
related characteristics of giant silkworm moth pupae; providing de- 
tailed descriptions of external morphology, length to width relation- 
ships for male and female antennae, and dimensional and weight data. 
The dimensional and weight data are of limited value, however, be- 
cause she mentions neither the number of pupae examined nor the 
sex. Mosher (1916a, Plates V & VI) also illustrates genital openings 
for a few species but does not discuss these structures. 

In some giant silkworm moths, such as Eupackardia calleta (West- 
wood) and Hyalophora cecropia (Linnaeus), pupae can be sexed cor- 


1 The opinions contained herein are those of the authors and should not be construed as official or reflecting the 
views of the Department of the Army. 

* Presently, Mathematical Statistician, USEPA Health Effects Research Laboratory, Research Triangle Park, North 
Carolina 27711. 

3 Presently, Associate Professor, Drinking Water Research Center, Florida International University, Miami, Flor- 
ida 33199. 


208 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


rectly in almost every instance because of consistently distinct dif- 
ferences in both the size of the antennae (Figs. 1-2) and the 
morphology of the genital openings (Figs. 5-6). In pupae of Callo- 
samia promethea (Drury) the dimensions of male and female anten- 
nae are not sufficiently different to permit reliable sex determinations 
(Figs. 3-4). Mosher (1916b) reported that the antennae of male C. pro- 
methea pupae are slightly longer and wider than those of the female, 
but that the antenna length is never more than three times the width 
in either sex. Also, the morphology of the genital openings can be 
highly variable in C. promethea. For certain groups of wild and reared 
pupae many individuals cannot be correctly sexed by examining these 
structures. The usual genital opening morphology for C. promethea 
pupae is shown in Figs. 7-8. 

Although there are observable differences in the size of male and 
female pupae of E. calleta, H. cecropia, and C. promethea, no infor- 
mation has been found in the literature to indicate that such differ- 
ences have been quantified or studied to determine their value in 
discriminating sex. Therefore, certain size characteristics of the pupae 
of these three species were examined to: (1) quantitatively define sex- 
related differences in weights and dimensions and; (2) determine sta- 
tistically whether such differences can be used singly, or in combi- 
nation, to discriminate sex. 


MATERIALS AND METHODS 


Measurements made during the study were: body weight (WT); 
body length (BL); body width (BW); circumference (CE); antenna 
length (AL); and antenna width (AW). Weights were determined to 
the nearest 0.01 gram using a Mettler H542 Analytical Balance. Mea- 
surements of BL, BW, and AW were made to the nearest millimeter 
using a vernier caliper. Body length was the distance from the vertex 
of the head to the posterior end of the abdomen; BW was the width 
at the 4th abdominal segment; and AW was the maximum width mea- 
sured perpendicular to the flagellum. Measurement of CE was made 
by placing a fine thread around the 4th abdominal segment and then 
measuring the thread on a metric ruler. Measurement of AW was 
made by placing a piece of monofilament nylon along the length of 
the flagellum and then measuring the piece of nylon on a metric ruler. 
Descriptive statistics (mean and 95% confidence interval) were cal- 


— 


Fics. 1-4. Ventral views of pupae. 1, male H. cecropia; 2, female H. cecropia; 3, 
male C. promethea; 4, female C. promethea. ANT, antenna; GO, genital opening; AO, 
anal opening. 


209 


VOLUME 36, NUMBER 3 


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210 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Fics. 5-8. Details of genital openings. 5, male H. cecropia; 6, female H. cecropia; 
7, male C. promethea; 8, female C. promethea. S8 and SQ, 8th and 9th abdominal 
segments; GO, genital opening; AO, anal opening. 


culated to quantitatively define sex-related differences. Data were 
subjected to discriminant function analysis (Freese, 1964) to deter- 
mine whether combination measurements, or ratios of measurements, 
were better discriminators of sex than individual measurements. Dis- 
criminant function analysis was used to estimate coefficients for a 
series of pupal measurements. Discrimination of sex is based on the 
magnitude of the discriminant function (Y) calculated from the for- 
mula Y = a,X, + a:X_ + agX3... + a,X,, where X; is a pupal measure- 
ment and a, is a coefficient. The method assumes for each type of 


VOLUME 36, NUMBER 3 LIL 


measurement that the variance is approximately the same for males 
and females. Generally, only those measurements that improve dis- 
crimination are used in the model. By using statistical tests (e.g., F) 
it is possible to determine, through sequential testing of pupal mea- 
surements, which coefficients differ from zero and should be included 
in the model. The sequential testing of coefficients assumes the nor- 
mal distribution. In addition to the single pupal measurements dis- 
cussed earlier, the following ratios were examined by discriminant 
function analysis: WT/AL, BL/AL, CE/AL, WT/AW, BL/AW, CE/AW, 
and AL/AW. Sources of specimens from which data were collected 
were various reared and wild specimens as follows: C. promethea 
were diapausing pupae collected in Harford County, Maryland, dur- 
ing the winter of 1973-74 (n = 34) and diapausing pupae collected 
in Portage County, Ohio, during the winter of 1974-75 (n = 43); E. 
calleta were diapausing pupae collected in various south Texas coun- 
ties during the fall of 1974 (n = 48); H. cecropia were diapausing 
pupae reared in Harford County, Maryland, during the summer of 
1973 (n = 65). Pupae that were later used to test the various discrim- 
inant function equations were either specimens reared in Frederick 
County, Maryland, or wild specimens collected in Frederick and Har- 
ford counties, Maryland, or various counties in south Texas. 


RESULTS AND DISCUSSION 


Pupal weights and dimensions for C. promethea, E. calleta, and H. 
cecropia are summarized in Table 1. Within species the sample means 
for males and females are significantly different (P < 0.05) for all mea- 
surements, except antenna length in C. promethea. The statistics 
shown in Table 1 are sample means, +95 percent confidence inter- 
vals: they define an interval for population means but not individuals 
in the population. Thus, the interval statistics characterize the values 
of male and female pupae of these three species but will not validly 
discriminate sexes in individual pupae. 

Discriminant function equations derived for determining sex in in- 
dividual pupae of the three species are shown in Table 2. The dis- 
criminant function analysis program derived numerous equations; 
those shown in Table 2 are judged the most predictive calculation for 
each species. The validity of these equations was tested by using 
them to predict sexes in groups of wild and reared pupae. For E. 
calleta, we collected the appropriate data from individuals in two 
groups of reared pupae and one group of wild pupae from south Texas. 
The discriminant function equation (DF,, = —10.19WT + 39.22AW) 
correctly calculated that there were 16 males and 16 females, and 7 
males and 12 females, respectively, in the two groups of reared pupae, 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


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VOLUME 36, NUMBER 3 | 213 


TABLE 2. Discriminant functions for determining the sex of giant silkworm moth 
pupae. 


Discriminant 
function equation Decision value 


Callosamia promethea: 


DF, = —2.69BW + 11.42AW female < 9.97 > male 

DF,, = 3.66(CE/AL) + 5.35(BL/AW) male < 49.33 > female 
Eupackardia calleta: 

DF,, = —10.19WT + 39.22AW female < 135.60 > male 
Hyalophora cecropia: 

DFy. = —2.25CE + 33.08AW female < 90.28 > male 


as confirmed by adult sexes at emergence. Similarly, 11 wild pupae 
of E. calleta were correctly sexed as 5 males and 6 females. For H. 
cecropia we examined an additional 11 reared pupae. This group con- 
tained 7 males and 4 females, as calculated by the discriminant func- 
tion equation (DF). = —2.25CE + 33.08AW) and confirmed at the time 
of adult emergence. For C. promethea we examined two additional 
groups of pupae containing 16 and 10 reared individuals, respective- 
ly; and a third group containing 33 wild individuals. The discriminant 
function equation (DF, = —2.69BW + 11.42AW) misclassified 2 fe- 
males in the first reared group, 7 females in the second reared group, 
and 5 females in the wild group, as determined by adult sexes at 
emergence. These misclassifications appeared to be due to a shift in 
the mean of some measurement used in the equation. Therefore, we 
used a similarly established predictive equation containing ratio in- 
formation (DF ,, = 3.66(CE/AW) + 5.35(BL/AW)) to determine wheth- 
er ratios might remain relatively constant when mean values were 
shifting. Using this second equation to calculate individual sexes of 
these three additional groups of C. promethea pupae resulted in sim- 
ilar misclassifications. 

These studies have quantified differences in the weights and di- 
mensions of male and female pupae of C. promethea, E. calleta, and 
H. cecropia. Antenna length in C. promethea was the only character- 
istic that was not significantly sex-related. This may account for the 
fact that antenna size in male and female individuals of this species 
is not a good discriminator of sex. Mosher (1916b) discusses antenna 
dimensions in terms of length to width ratios, but does not present 
data on absolute sizes. There is no known published information on 
the origin of describing lepidopterous antennae using ratios. The 
characteristic actually being perceived by an observer is the antenna 
surface area, and an approximation of that characteristic would seem 


214 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


ANTENNA 
LENGTH: WIDTH RATIO 


——_ 75 


50- 


ANTENNA SURFACE AREA (mm? 
i.e) 
on 


cP EC HC cP EC HC 


Fics. 9-10. Antenna size relationships in pupae of C. promethea (CP), E. calleta 
(EC), and H. cecropia (HC). 9, comparative length to width ratios; 10, comparative 
surface areas. 


VOLUME 36, NUMBER 3 215 


to be more valuable than the length to width ratio. The antenna sur- 
face area (SA) can be approximated (SA = 0.5(AL x AW)) from the 
length to width information. Antenna length to width ratios and an- 
tenna surface areas are compared for male and female pupae in Figs. 
9-10. The comparison quantifies the fact that there is less visually- 
perceptible difference in male and female antennae in C. promethea 
than the other two species and thus, more difficulty in discriminating 
sex on the basis of antenna size in C. promethea. Another point ap- 
parent from the length to width information is that for each sex the 
ratios do not differ significantly among the three species. Whether 
these very similar ratios for each sex have a relationship to the sensory 
function of the antennae in the adult moths is not known. 


CONCLUSIONS 


In C. promethea neither the combination measurements nor the 
ratios of dimensions used in the discriminant function analysis re- 
sulted in reliable equations for determination of individual sexes. Ex- 
amination of the genital openings appears to be the best available 
way of determining sex in individual C. promethea pupae. 

For E. calleta pupae, weight and antenna width were reliable in- 
dicators of individual sex when used in the discriminant function 
equation derived for this species. 

For H. cecropia pupae, discriminant function analysis demonstrat- 
ed that circumference and antenna width data are the most reliable 
dimensions for discriminating sex in individuals. 


ACKNOWLEDGMENT 


We acknowledge the assistance of R. S. Peigler in providing wild Eupackardia cal- 
leta (Westwood) pupae from south Texas; with these we were able to verify the dis- 
criminant function equation for that species. 


LITERATURE CITED 


Butt, B. A. & E. CANTU. 1962. Sex determination of lepidopterous pupae. U.S. Dept. 
Agri., ARS 33-75. 7 pp. 

EHRLICK, F. T., R. T. FRANKLIN & R. N. COULSON. 1969. Characters for determining 
sex of pupae of the oakworms, Anisota senatoria, A. stigma, and A. virginiensis, 
and the yellow-neck caterpillar, Datana ministra. Ann. Entomol. Soc. Amer. 62(4): 
931-932. 

FREESE, F. 1964. Linear regression methods for forest research. U.S. Forest Service 
Research Paper, FPL 17. December 1964. 

JENNINGS, D. T. 1974. Sexing southwestern pine tip moth pupae, Rhyacionia neo- 
mexicana (Lepidoptera: Olethreutidae). Ann. Entomol. Soc. Amer. 67(1):142-143. 

KEAN, P. J. & A. P. PLATT. 1973. Methods for externally sexing mature larvae and 
pupae of Limenitus (Nymphalidae). J. Lepid. Soc. 27(2):122-129. 

MOSHER, FE. 1914. The classification of the pupae of the Ceratocampidae and Hemi- 
leucidae. Ann. Entomol. Soc. Amer. 7(4):277-300. 


216 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


1916a. A classification of the lepidoptera based on characters of the pupae. 

Bull. Ill. State Lab. Nat. Hist. 12(2):13-159. 

1916b. The classification of the pupae of the Saturniidae. Ann. Entomol. Soc. 
Amer. 9(2): 136-158. ; 

MuccLul, J. M. 1974. Sex identification of Malacosoma disstria (Lepidoptera: Lasio- 
campidae). Ann. Entomol. Soc. Amer. 67(3):521-522. 

SOLOMON, J. D. 1962. Characters for determining sex in elm spanworm pupae. J. 
Econ. Entomol. 55(2): 169-170. 

VILLIARD, P. 1969. Moths and how to rear them. Funk and Wagnalls, New York, New 
York. 242 pp. 


Journal of the Lepidopterists’ Society 
36(3), 1982, 216-217 


BOOK REVIEW 


MICROLEPIDOPTERA, by Elwood C. Zimmerman. 1978. Insects of Hawaii, volume 9, 
xxiv + 1903 pages, 1355 cuts. Price: U.S. $60. University of Hawaii Press, Honolulu. 


The first 200 pages are an overview of the Lepidoptera that includes a) classification, 
b) identification keys to the immature stages of the species found in Hawaii and sep- 
arate keys to the larvae and pupae found in specific habitats, c) morphology of the 
immatures and adults, and d) techniques for preparing and handling adults and im- 
matures for collections and for wing and genital studies. A checklist of the described 
(only previously described or misidentified species are treated) genera and species and 
list of nomenclatural changes are useful. Fourteen of the 80 genera and 605 of the 681 
species are endemic. A synopsis of the distribution of genera and species by island 
illustrates the high degree of species’ endemicity; however, lack of adequate collecting 
makes the tables preliminary. 

The systematic treatment is a good survey of the microlepidoptera of Hawaii. Ilus- 
trations are abundant and cover the immature as well as the adult stages. Zimmerman 
has developed identification keys to most of the taxa. A notable exception is to the 
species of Hyposmocoma Butler. New generic names are proposed and defined when 
necessary. He has brought together published drawings of structural parts of adults 
and larvae of as many species as possible and reproduced them in this volume; so, the 
user has before him much of the extant illustrative material. 

Zimmerman proposes a relatively conservative higher classification at the family 
level within the so-called microlepidoptera that is particularly noteworthy in the Ge- 
lechioidea. He places the Oecophorinae, Ethmiinae, Xyloryctinae, Blastobasinae, Chry- 
sopeleiinae, Momphinae, Cosmopteriginae, and Gelechiinae as subfamilies of the Ge- 
lechiidae. Based on strict priority of family-group names, the superfamily and family 
should be Oecophoroidea and Oecophoridae (Bruand, 1850), not Gelechioidea and 
Gelechiidae (Stainton, 1854). I agree with Zimmerman’s philosophy on the inflation of 
the classification of the microlepidoptera but not with all of his conclusions. However, 
the final word definitely is not written on classification, particularly that of the Gele- 
chioidea. Some major differences are the following: Thyrocopa Meyrick is in the Au- 
tostichinae of the Oecophoridae rather than in the Xyloryctinae of the Gelechiidae. 
Chedra Hodges and Batrachedroides Zimmerman are in the Batrachedrinae of the 
Coleophoridae rather than in the Momphinae, Gelechiidae. Momphidae, sensu stricto, 
do not occur in Hawaii. Cosmopterigidae have two subfamilies in Hawaii, Cosmop- 
teriginae with four genera, and Chrysopeleiinae with one introduced species and ge- 
nus. Symmocinae are a subfamily of Blastobasidae rather than a tribe of the Gelechi- 
inae. The correct spelling for Dichomerini is Dichomeridini. Sitotroga Heinemann 
and Pectinophora Busck are in the Chelariinae rather than the Gelechiinae. Merim- 
netria Walsingham is in the Anomologinae rather than the Aristoteliini. 


VOLUME 36, NUMBER 3 ley 


Defining the limits of genera is often one of the most difficult problems that taxon- 
omists face. Zimmerman’s perplexion in dealing with the extremely large genus Hy- 
posmocoma, with an estimated 500+ species endemic to Hawaii, is understandable. 
He synonymized 13 generic names under Hyposmocoma after finding no consistent 
differences to separate groups associated with the previously proposed generic names. 
Six hundred thirty pages of the two-volume work are devoted to the 350 described 
species of Hyposmocoma. | strongly recommend this section as an example of the 
morphological diversity that can occur in a genus. Zimmerman states that a life’s work 
could be spent on this genus. With a projected 150 undescribed species and without 
an identification key to the described species, determination of species is problemat- 
ical, but a far greater start has been made to deal with them than ever before. He lists 
a number of criteria that he discovered to be significant and that may be helpful to the 
person who attempts to devise such a key. 

The text is written in a highly readable style. It is often interspersed with biting 
comments relative to lack of support for this immense project that, without personal 
knowledge of his situation, may appear incongruous. I strongly agree with his often- 
made observation that Hawaii has been visited by many, but our knowledge of the 
microlepidoptera that occur on the islands is so inadequate as to be appalling—the 
same is true for the continental United States. From the vantage point of a systematist 
working at the U.S. National Museum of Natural History and considering that Hawaii 
is one of the States, I am a bit troubled that nearly all of the specimens reside in the 
British Museum of Natural History. 

By and large typographic errors are few. The most annoying feature of the book is 
that often the text is cut up and separated by illustrations, in one instance with 124 
pages of them. But, I can understand the view of wanting to have the illustrations near 
the associated text. I am puzzled by the large number of illustrations of adult mor- 
phology that do not accompany any text. The wealth of information about Lepidoptera 
in general and the microlepidoptera specifically is welcomed. All students of the Hawai- 
ian biota and of the microlepidoptera will find much of interest, many facts to digest, 
and many leads for further research. I congratulate Zimmerman on a fine accomplish- 
ment. 


RONALD W. HODGES, Systematic Entomology Laboratory, IIBII, SEA-AR, USDA, 
Washington, D.C. 20560. 


Journal of the Lepidopterists’ Society 
36(3), 1982, 218-226 


CLASSIFICATION AND LIFE HISTORY OF ACSALA ANOMALA 
(ARCTIIDAE: LITHOSIINAE) 


J. DONALD LAFONTAINE 


Biosystematics Research Institute, Research Branch, Agriculture, 
Canada, Ottawa, Ontario KIA 0C6 


JOHN G. FRANCLEMONT 


Department of Entomology, Comell University, 
Ithaca, New York 14853 


AND 


DOUGLAS C. FERGUSON 


Systematic Entomology Laboratory, U.S. Dept. of Agriculture, U.S. National Museum, 
Smithsonian Institution, Washington, D.C. 20560 


ABSTRACT. The immature stages and female of Acsala anomala Benjamin are 
described for the first time from material collected in the Yukon Territory, Canada, 
during 1980. The position of the genus within the Noctuoidea is reassessed on the 
basis of adult and larval characters. It is concluded that the genus Acsala should be 
transferred from the Lymantriidae to the subfamily Lithosiinae in the Arctiidae. The 
life history of Acsala anomala is described and the immature stages illustrated. 


In 1980 J.D.L. had the opportunity to collect in the Yukon Territory 
with Dr. D. M. Wood, also of the Biosystematics Research Institute. 
The purpose of the trip was to initiate a study of the insects and 
arachnids of northwestern North America, particularly those of the 
unglaciated Beringian refugium area. 

During the summer, Lafontaine and Wood devoted a considerable 
amount of time to a study of the immature stages of the little known 
moth Acsala anomala Benjamin. This was done because of uncer- 
tainty of the phylogenetic position of Acsala within the Noctuoidea 
based on characters of the adult male and because of interest in its 
classification by J.G.F. and D.C.F. Previous to 1980 the species was 
known from less than a dozen specimens, all males. Collections of 
Acsala anomala were made in the northern Ogilvie Mountains in 
north-central Yukon and in the Richardson Mountains in northern 
Yukon. Adults of both sexes and immature stages were collected and 
their habits were recorded in the field. The following discussions on 
the classification and life history of Acsala anomala are based on these 
specimens and field data. | 


CLASSIFICATION 


Benjamin (1935) in describing Acsala put it in the Arctiidae, al- 
though he suggested that it should probably be in a separate family. 


VOLUME 36, NUMBER 3 219 


Benjamin believed that Acsaia was a remnant of a primitive group 


that gave rise to the Arctiidae and Lymantriidae. McDunnough (1938) 
put Acsala in the subfamily Arctiinae next to Dodia albertae Dyar, 
presumably because of convergent similarities resulting from similar 
adaptations to an arctic environment (e.g., dark coloration; hairy, 
translucent wings; aborted proboscis). Ferguson (1978), however, 
showed that structural characters do not support classification of Ac- 
sala within the Arctiinae. Lack of ocelli, a character of Acsala, is a 
distinguishing feature of the arctiid subfamily Lithosiinae and of the 
Lymantriidae. Wing venation of Acsala could be either arctiid or 
lymantriid. It has a long discal cell, sometimes regarded as a charac- 
ter of the Lithosiinae and of other Arctiidae, but various exotic 
Lymantriidae (e.g., species of Eloria Walker) have discal cells that 
are as long or longer. Characters more clearly suggesting an affinity 
with the lithosiines are: presence of an orange prothoracic collar 
and lack of long terminal spinules on the male antenna; the female 
antenna is filiform. Ferguson classified Acsala within the Lymantri- 
idae on the basis of venation, especially the relationship of veins 
Sc and R in the hind wing, which is unusual for an arctiid; however, 
he expressed considerable reservation, pointing out that characters 
of the larva were needed in order to resolve the problem. 

The discovery of the larva makes possible a reassessment of the 
position of Acsala; the larval characters resolve the dilemma, “Arc- 
tiidae or Lymantriidae?,”’ that so troubled Benjamin (1935) and Fer- 
guson (1978). The larva is a lithosiine arctiid. The general appearance, 
at first glance, is that of an ordinary arctiid (Fig. 5); the setae are 
barbed and arise from verrucae. It differs from known lymantriid lar- 
vae in that there are no dorsal glands on abdominal segments six and 
seven. The distinctive character that it shares with other known lith- 
osiine larvae is the possession of a conspicuous mola on the inner 
face of the mandible at the base (Fig. 15). The mandible of a member 
of the Arctiinae (Fig. 16) is illustrated for comparison. Gardiner (1943) 
seems to have been the first to notice this character and to have stressed 
its importance in characterizing the lithosiines. Acsala larvae, like 
those of some other lithosiines, e.g., Eilema species and Lithosia 
quadra (Linnaeus), have the crochets heteroideous (Fig. 13) as do the 
larvae of the Arctiinae and Ctenuchinae. It must be noted that many 
Lithosiinae, e.g., Asura anomala (Elwes), A. calamaria (Moore), 
Chionaema bianca (Walker), C. detrita (Walker), C. peregrina (Walk- 
er), Hypoprepia miniata (Kirby), Clemensia albata Packard, Eudes- 
mia species, Crambidia “white species,’ and Agylla species, have 
the larval crochets homoideous. However, in all these larvae the man- 


220 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


“Me 


> 


CR show 


ear 
a 


Fics. 1-6. Acsala anomala. 1, habitat—Km 465 Dempster Hwy., Yukon Territory; 
2, larva feeding, 1.5x; 3, larvae resting under rock, 0.6x; 4, third instar larva, 2.5x; 
5, mature larva, 4x; 6, adult female, 3x. (Photographs by J. D. Lafontaine and Bio- 
graphic Unit, Agriculture Canada.) 


— 


Fics. 7-12. Acsala anomala. 7, adult male, 1x; 8, female laying eggs, 1.7x; 9, 
lateral view of egg, 40x; 10, dorsal view of egg, 40x; 11, details of egg microsculpture, 
80x; 12, micropylar region at apex of egg, 400. (Photographs by J. D. Lafontaine and 
Bio-graphic Unit, Agriculture Canada.) 


221 


VOLUME 36, NUMBER 3 


UPA JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


dible has a basal mola. The larval foodplants, various species of li- 
chens, are also typical of lithosiine arctiids. 

The first instar larva of Acsala has single, barbed setae arising from 
inconspicuous, raised pinacula; there are two prespiracular setae on 
T,, and each of the thoracic segments has two setae, group vii or vi, 
above the leg. 

In the last larval instar T, has a moderately large prespiracular ver- 
ruca with many setae. T; has one dorsal verruca, that is there are two 
verrucae above the prespiracular line and two below the prespiracular 
line and above the leg. A; has three verrucae above the spiracle, a 
small verruca just behind and slightly below the spiracle, and two 
large verrucae below the spiracle and above the proleg, the latter with 
numerous secondary setae. There is no indication of viii on the four 
proleg bearing segments. Verruca pattern on A; is similar to that shown 
for Eilema sp. by Gardiner (1943, Fig. 9) but the small verruca behind 
the spiracle has several setae. A, and A, as in A;, but with small 
verrucae at the vii and viii positions. A, and Ag, similar to A, and Ag. 

On the basis of our analysis of larval characters, the genus Acsala 
was transferred in the North American Lepidoptera check list from 
the Lymantriidae to the subfamily Lithosiinae in the Arctiidae (Fran- 
clemont, in press). 

In the adult only the venation seems to warrant consideration be- 
cause Benjamin and Ferguson discussed other characters so thor- 
oughly. Considerable emphasis has been placed on the relative po- 
sition of Sc and R of the hind wing in the Noctuoidea. However, it 
becomes evident when many species of the various families are stud- 
ied that this relationship is more variable than keys and characteriza- 
tions of the venation would imply. In the lithosiines the relationship 
is highly variable; the base of R is missing, or fused with Sc, in most 
lithosiines but in the genus Hypoprepia the base of R is present but 
weak. In Acsala anomala the venation is variable within the species. | 
In 23 of 40 specimens examined, Sc makes a sharp bend down to R, 
touches, and then separates immediately before the middle of the 
discal cell; in nine specimens the relationship is much as described 
_ by Benjamin, Sc and R are connected by a bar. The drawing in Fer- 
guson (1978) shows Sc with a sharp bend, but it does not touch R; 
this condition occurs in eight specimens. Except for the relationship 
of Sc and R, the hind wing venation of Acsala is similar to that of the 
lithosiine Clemensia albata Packard; in most specimens R and M, of 
the hind wing are long-stalked, separating a short distance before the 
wing margin. In one specimen vein R is absent and in another vein 
M; is missing. The forewings have no accessory cells in about one 
quarter of the specimens. Many specimens show differences in length 


VOLUME 36, NUMBER 3 Dae 


of stalking and in size of the accessory cell in the wings on the op- 
posite sides of the same moth. Such variation in wing venation may 
be related to a tendency toward flightlessness in both sexes. The fe- 
male is flightless (Fig. 6) and wing size in males is variable. The wing 
venation of the female is similar to that of the male but the veins are 
crowded together because of smaller size. 


Interpretation of Characters of Acsala 


Although many of the characters of Acsala are typical of the Arcti- 
idae, most of them cannot be used, at present, for phylogenetic inter- 
pretation. The absence of dorsal glands on abdominal segments six 
and seven is a character state of the larvae that is primitive in the 
Noctuoidea and cannot be used for phylogenetic interpretation within 
the superfamily. Other character states of Acsala, the configuration of 
larval verrucae, male antennae characters, and adult wing venation 
characters, may be derived in the Arctiidae and phylogenetically sig- 
nificant. Assessment of these characters will not be possible until the 
relationship of the Arctiidae and the Lymantriidae has been deter- 
mined. Two definitive character states of Acsala that allow it to be 
placed in the lithosiine arctiids are the heteroideous crochets of the 
larva (Fig. 13) and the presence of a mola on the inner face of the 
mandible (Fig. 15). These derived character states are restricted to 
the Arctiidae and the Lithosiinae respectively. The absence of ocelli 
is more difficult to interpret because their loss may have occurred a 
number of times in the Noctuoidea. 


LIFE HISTORY 


When Lafontaine and Wood arrived in the Ogilvie Mountains in 
north-central Yukon on 18 June, males were already in flight. The 
flight season lasted until late June at low elevations (800-1000 me- 
ters), but adults were collected as late as 21 July at high elevations 
(1600 meters). Males fly over loose, rocky slopes and hilltops where 
vegetation is sparse (Fig. 1). Butterflies commonly found in associa- 
tion with it are Erebia magdalena mackinleyensis Gunder and Bo- 
loria (Clossiana) astarte distincta (Gibson). The flight of the moth is 
weak and fluttering, similar to that of a large caddisfly. In spite of this, 
they are hard to catch because of difficult footing on the rocky slopes 
and because males tend to move along rapidly in the wind. Most 
activity is confined to bright, sunny periods. Males rest during cloudy 
periods and at night when the sun is low on the horizon. 

After collecting a series of males, an attempt was made to find a 
female. Males were followed in the hope that they would go to a 
female; however, after several days none was found. In late June we 


224 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


~ 


~ 


cae 
a ‘ 


\ 


a 
-~ 


~ 


au 


WY 


WY 
WAN Ae TREN NAN TS 
a nan Ls AS 


Pi} PAT ENS 


AS 


13 


Fics. 13-16. Details of larval characters of Acsala anomala. 13, inside view of Ag 
proleg showing heteroideous crochets; 14, lateral view of A;; 15, inner surface of man- 
dible showing mola; 16, inner surface of mandible of Arctiinae, Spilosoma vagans 


(Boisduval) for comparison. 


VOLUME 36, NUMBER 3 225 


traveled north to the Richardson Mountains where eggs, larvae, empty 
cocoons, and several females were found by searching under rocks 
on rocky slopes and hilltops. Females spend most of the time under 
rocks but probably crawl onto the upper surface to mate. One female 
that had just mated was observed on top of a rock with a male. Egg 
batches of 6 to 30 eggs are laid in a single layer on the underside of 
a rock. Eggs are reddish orange when laid. They turn pale orange 
after about a week. Several hours before hatching, the black larval 
head capsule is visible through the eggshell. Details of egg sculptur- 
ing are shown in Figs. 9-12. 

Larvae hatch from the eggs in eight to ten days, eat their eggshell, 
and then begin to feed on lichens. The first instar larva has a yellow- 
orange body with a black head and is about 2 mm long. The larvae 
hide during the day (Fig. 3); they feed on lichens during the evening 
and at night when the sun is lower (Fig. 2). The larvae apparently 
feed almost exclusively on black colored, low, foliose lichens (Para- 
melia stygia (L.) Ach. and Umbilicaria cf. proboscidea (L.) Schrad.) 
and on black crustose lichens (Orphniospora atrata (Sm.) Poelt, Buel- 
lia cf. spuria (Schaer.) Anzi, Lecidia armeniaca (DC.) Fr., and L. 
fuscocinerea Ny]l.). 

The larvae probably take many years to mature. In June when adults 
are in flight, eggs and larvae of all instars except the first were found. 
The locations of several egg batches were marked in June; by the end 
of the summer most larvae had just molted to second instar. The larvae 
may tend to crawl upwards; abundance of larvae increases moving up 
a hill with the greatest densities occurring at the hilltop. Egg batches 
and females, however, were lower down, on the hillsides. The hilltop 
shown in Fig. 1 had larvae under almost every rock. 

Cocoons are spun and attached to the underside of a rock. Larval 
hair is used in its construction. The pupa lacks excessive body hair 
found in lymantriid pupae but does have small clusters of hair at 
verrucae scars as is typical of arctiid pupae (Mosher, 1914). We could 
not determine whether mature larvae pupate in the fall or in the spring. 
When we left the Richardson Mountains at the end of the season, 
mature larvae but no pupae were found. After bringing them into the 
laboratory in late August several did pupate and adults emerged by 
mid-September. Larvae were relatively easy to rear on location where 
fresh lichens could be supplied each day. Rearing was difficult under 
laboratory conditions, however, because the lichens tended to dry out 
and if moistened, they would mildew. 


Distribution and Abundance 


Acsala anomala is known only from unglaciated areas in the north- 
ern half of Yukon and Alaska. It is probable that its range remained 


226 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


restricted following deglaciation because of habitat limitations and 
poor dispersal ability. 

At first it seemed puzzling that a species that is so abundant as 
larvae should be relatively scarce as an adult. This anomaly can be 
explained by two factors; first, if the larvae take a number of years to 
mature, only a fraction of the larvae would pupate each year. Second, 
parasitism may take a heavy toll. Cocoons of a braconid parasite, Me- 
teorus sp., were more common than were those of the moth. A tach- 
inid parasite, Tryphera sp., was also present. In addition to these, an 
ichneumonid hyperparasite, Gelis obesus (Ashmead), was reared, this 
apparently attacking a second, uncollected, braconid parasite. 


ACKNOWLEDGMENTS 


We thank T. L. McCabe, New York State Museum, Albany, and J. E. Rawlins, Cornell 
University, Ithaca, New York, for furnishing unpublished data on the condition of the 
crochets in some species of lithosiines. We also thank J. R. Barron, W. R. M. Mason 
and D. M. Wood, Biosystematics Research Institute, Ottawa, for identifying ichneu- 
monid, braconid and tachinid parasites, and I. M. Brodo, Botany Division, National 
Museums of Canada, Ottawa for identifying the lichens. We thank J. A. Downes and 
D. M. Wood, B.R.I., Ottawa, for reading and commenting on the manuscript. 


LITERATURE CITED 


BENJAMIN, F. H. 1935. A new genus and species from Alaska (Lepid., Arctiidae, 
Nyctemerinae). Can. Entomol. 67:195—197. 

FERGUSON, D. C. in R. B. DOMINICK et al. 1978. The moths of America north of 
Mexico, Fasc. 22.2, Noctuoidea (in part): Lymantriidae. E. W. Classey Limited and 
The Wedge Entomological Research Foundation, London. _ 

FRANCLEMONT, J. G. Arctiidae. In R. W. Hodges, F. M. Brown, D. R. Davis, D. C. 
Ferguson, J. G. Franclemont, J. B. Heppner, L. D. Miller, E. G. Munroe and 
E. L. Todd. Check list of the Lepidoptera of America north of Mexico. E. W. 
Classey Limited and The Wedge Entomological Research Foundation, London (in 
press). 

GARDINER, J. C. M. 1943. Immature stages of Indian Lepidoptera (5). Indian J. Ento- 
mol. 5:89-102. 

McDUNNOUGH, J. 1938. Check list of the Lepidoptera of Canada and the United 
States of America. Part 1, Macrolepidoptera. Mem. Sth. Calif. Acad. Sci. Vol. 1. 

MOSHER, E. 1916. A classification of the Lepidoptera based on characters of the pupa. 
Bull. Il. St. Nat. Hist. Surv. 12:17-160. 


Journal of the Lepidopterists’ Society 
36(3), 1982, 227-228 


GENERAL NOTES 
A BILATERAL GYNANDROMORPH OF ERYNNIS HORATIUS (HESPERIIDAE) 


Eight larvae (2nd and last instars) and 2 pupae of Horace’s dusky wing, Erynnis 
horatius Scudder and Burgess, were collected on 13 August 1980 in Baton Rouge, East 
Baton Rouge Parish, Louisiana. The larvae were found on cherrybark oak, Quercus 
falcata var. pagodaefolia Ell. and raised on this in captivity. On 10 September 1980 
an adult bilateral gynandromorph emerged. 

The right half of the specimen is female in appearance, and the left is male (Fig. 1). 
The right forewing measures 18.1 mm in length and exhibits characters of a typical 
phenotype 2 female, i.e., spots 2, 3, 6, 7, 8, 9 and Yd 5 are present (Burns, J. M., 1964, 
Evolution in skipper butterflies of the genus Erynnis, Berkeley, Univ. of Cal.). The 
left forewing measures 16.4 mm in length and has the male costal fold, spots Yd 5 and 
3 are smaller in the left forewing (¢) than in the right forewing (2), and spot 2 is not 
present on the left forewing (¢). The median dark brown mottling on the right (2) 
forewing is stronger than on the left (¢) forewing and the right hindwing (¢) is also 
lighter than the left (¢) hindwing. All spots on both forewings are semihyaline. The 
antennae appear dissimilar and the right (2) labial palp measures 1.5 mm and left (d) 
1.2 mm. 

The external genitalia were dissected and consisted of male structures including an 
aedeagus. A bursa copulatrix was also present. The valvae of a typical individual (com- 


Fic. 1. Bilateral gynandromorph of Erynnis horatius Scudder and Burgess. 


228 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Fic. 2. Left and right valvae of Erynnis horatius a) typical individual, b) corre- 
sponding gynandromorph. 


pared with illustrations from Forbes, W. T. M., 1960, Lepidoptera of New York and 
neighboring states, Part IV, Cornell Univ. Agri. Exp. St. Memoir 371) and the corre- 
sponding pair of gynandromorph valvae are shown in Fig. 2. The length of the right 
and left valvae of the typical individual (collected 23 August 1976, Baton Rouge, LA) 
measures 1.15 mm and 1.30 mm respectively, while the gynandromorph valvae mea- 
sure 0.95 mm and 1.05 mm. The gynandromorph has ampullae on both valvae, while 
the typical individual displays the usual asymmetrical dimorphism. The number of 
marginal spines on the valvae of the typical specimen is 11 (right) and 11 (left) in 
contrast to that of the gynandromorph which is 13 (right) and 9 (left). 

The specimen and pupal case have been deposited in the Louisiana State University, 
Department of Entomology Collection (LSUC). We thank Drs. H. B. Boudreaux and J. 
B. Chapin for reviewing the manuscript. 


MICHAEL L. ISRAEL & JAMES E. CILEK, Department of Entomology, Louisiana Ag- 
ricultural Experiment Station, Louisiana State University, Baton Rouge, Louisiana 
70803. 


Journal of the Lepidopterists’ Society 
36(3), 1982, 229-230 


THE CASE OF PERRHYBRIS LYPERA (PIERIDAE) AND THE 
LAURACEAE: HOST-PLANT RECORD OR ASSUMPTION? 


The study on Perrhybris lypera (Pieridae) in Costa Rica by Young (1980, J. Lepid. 
Soc. 34:36-47) reveals some interesting differences from my own observations on that 
species and raises some points I would like to clarify concerning host-plant records. 
While engaged in studies on the butterflies of the La Selva field station, Heredia 
Province, Costa Rica, I was able to study P. lypera intermittently for a period of six 
months (July through December 1979). Both study sites used by Dr. Young (La Selva 
and “La Tigra’) are part of the same tract of forest and have adjoining boundaries. 
Although “La Tigra” is under extensive cultivation, the remnant patches of forest there 
have a considerable number of both butterfly and plant species in common with La 
Selva. 

While it is of great interest that Dr. Young has found P. lypera in association with 
Ocotea sp. (Lauraceae), because it represents a divergence from host-plant relation- 
ships known in the Pieridae, my own experiences with P. lypera at La Selva differ 
considerably, and I feel that he is in error concerning his host-plant record. I have in 
eleven instances while at La Selva reared P. lypera to adulthood and all were on 
Capparis pittieri (Capparidaceae). This is in accord with my other host-plant records 
for other Perrhybris species in Costa Rica and Peru as well as records of other workers 
(L. Gilbert, M. Singer & J. Smiley, pers. comm.). These rearings are also in accord with 
my host-plant records for other closely related genera (Itaballia and Pieriballia) in 
Costa Rica, which likewise feed on Capparidaceae as larvae. While Dr. Young has 
observed an oviposition record by P. lypera on the host-plant (which I believe is 
Capparis) and obtained first instar larvae, he has not reared them to adulthood. This 
does not constitute a host-plant record for the butterfly; reared butterflies are from 
larvae that feed on and develop to adulthood on a certain plant. In figure 2 (Young, 
op. cit.) several photographs are shown with eggs on the upperside of a leaf along 
with a photograph of the first instar larvae. All of the photographs show “pronounced 
stellate pubescence,” which appears to me to be highly characteristic of many Cap- 
paridaceae in Costa Rica, yet I know of no Lauraceae occurring at La Selva (Ocotea 
in particular) which show this character. It is my suspicion that Dr. Young has con- 
fused the identity of the larval host-plant with that of the leaf he originally found 
the pupae of P. lypera upon. I therefore question his speculations on the aposematic 
nature of P. lypera, because they are based on suspect host-plant data, not because 
of features inherent to the natural history of the butterfly. 

His assumption that the pupation site constitutes the larval host-plant may be mis- 
leading. In Dr. Young’s paper the assumption is made that the genus Pereute (Pieridae) 
uses Ocotea, and this was perhaps influential in his recording P. lypera as the second 
record of a member of the Pieridae to feed on the Lauraceae; both are very interesting 
records. This assumption is based on Jorgenson (1916, Ann. Museo Nacional, Buenos 
Aires 28:427-520), which says that groups of larvae and pupae of Pereute were found 
on the trunk of an Ocotea tree. However, my own field work in Costa Rica indicates 
that the genus Pereute does not feed on the Lauraceae as has been assumed. Pereute 
and other closely related genera in Costa Rica feed on the Loranthaceae (DeVries, ms. 
in prep.), which are common epiphytic parasites of many tropical forest trees, including 
Ocotea. Larvae feeding on these epiphytes craw] down the tree and pupate on the 
tree trunk. They do not feed on the leaves of the plant where pupation takes place. 
While the use of the Loranthaceae by New World Pieridae is still somewhat novel (i.e., 
unstudied), the allied genus Delias in the Old World uses Loranthaceae extensively, 
and their pupation behaviors are similar to those of Pereute; both genera follow the 
theoretical lines of coevolution of butterflies and plants of taxonomic relatedness. Thus 
the genera Pereute and Perrhybris both appear to be erroneous records on Lauraceae 
and have little in common regarding their respective host-plants. 

As tempting as it may be, unless larvae actually feed upon and develop into adults 


230 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


on a particular plant, I do not feel one can draw lines of host-plant relationships by 
casual observation. With this in mind I would urge future workers to be suspicious of 
host-plant records that are far afield from what we know of coevolutionary relationships 
(Ehrlich & Raven, 1965, Evolution 18:586-618), to double check the identity of host- 
plant material that is taken from the plant on which larvae are feeding, and to record 
oviposition observations as such. By following such criteria, perhaps future miscon- 
ceptions and errors can be minimized. 


PHILIP JAMES DEVRIES, Department of Zoology, University of Texas, Austin, Texas 
USE MD, 


Journal of the Lepidopterists’ Society 
36(3), 1982, 230-232 


PERRHYBRIS LYPERA (PIERIDAE) FEEDING ON LAURACEAE: 
A RESPONSE TO DEVRIES 


In an earlier paper in the pages of this journal (Young, 1980, J. Lepid. Soc. 34:36- 
47) I reported both oviposition and first instar larval feeding on the young leaves of a 
tropical rain forest understory tree identified, albeit from vegetative parts alone, by 
reputable authorities as a species of Ocotea in the Lauraceae. I had originally discov- 
ered the gregarious pupae of this butterfly on a mature leaf of a tree from Finca La 
Selva in 1969 and tentatively identified at that time by Dr. William Hathway of the 
University of Washington as either Ocotea or Nectandra (both Lauraceae). The sub- 
sequent observations, several years later, of oviposition and larval feeding at Finca La 
Tigra, approximately ten km from the La Selva site but at a slightly higher elevation, 
also revealed an association with Lauraceae (Young, op. cit.). 

Mostly by accident and indirect communication, I learned of the note by Mr. DeVries 
already submitted to this journal (DeVries, 1982, J. Lepid. Soc. 36:229-230), in which 
he suggested an error in the identification of the oviposition and larval host for P. 
lypera which I reported (Young, op. cit.). At my request, Mr. DeVries very graciously 
sent me a copy of his note. At the time I was preparing to leave for Costa Rica, and 
therefore, had the timely opportunity to once again check in the wild the food plant 
questioned. 

I retrieved additional samples of the leaves and stems of the exact same tree, a feat 
made simple because that tree had been marked for further studies of P. lypera be- 
havior and natural history at this locality. This fresh material was taken to San Jose 
where Dr. Gary Hartshorn, the well-known authority on tropical trees who identified 
Mr. DeVries’s La Selva food plant of P. lypera, made an identification of my material. 
Thus, the opportunity offered a control of sorts, since P. lypera food plant materials 
from two different sources (DeVries and Young) would have been identified by the 
same authority, something indeed worth doing if an error had been made by other 
authorities in earlier identifications. Dr. Hartshorn kindly examined my fresh material 
and gave me his very assured identification of the tree as Nectandra gentlei (Laura- 
ceae). He also indicated to me that, while the tree was very clearly lauraceous, the 


VOLUME 36, NUMBER 3 ell 


t 4 * 3 » ~ by) . : ° . ; 
2. ee = 2 ea = z i : . . aa ‘ad ee 


Fic. 1. Mature leaves of Nectandra gentlei (Lauraceae) from “Finca La Tigra’ in 
northeastern Costa Rica possess a thick coating of pubescence. N. gentlei is a larval 
food plant of the pierid Perrhybris lypera at this locality (see also Young, 1980, J. Lepid. 
Soc. 34:36—47). 


genera Ocotea and Nectandra are extremely closely related, and eventually, the former 
will probably be combined with the latter in a systematic revision. I explained the P. 
lypera debate to Dr. Hartshorn, who then commented to the effect that it is easy to 
distinguish between Capparidaceae and Lauraceae. Dr. Hartshorn thus confirmed my 
original identifications of this food plant as being Lauraceae, and I, therefore, conclude 
that an error in identification in Young (op. cit.) had not been made. Dr. Hartshorn 
agreed to deposit my pressed material of this tree in the herbarium collections of the 
National Museum of Costa Rica. 

DeVries (op. cit.) offers as proof of an error in identification the stellate pubescence 
characteristic of the young leaves fed upon by P. lypera in my study (my figure 2 in 
Young, op. cit.). He claims that Lauraceae do not have such a characteristic. Yet, it is 
common knowledge, particularly in dealing with tropical evergreen floras, that mor- 
phological characteristics of young leaves of a tree can be quite different from older 
leaves on the same tree. Mr. DeVries’s claim that the Lauraceae, including the food 
plant genus reported in my paper, do not possess such pubescence is not substantiated 
by available data. Fig. 1 shows the pubescence from a mature leaf of the larval food 
plant from La Tigra. This pubescence, while not as pronounced as in the young leaves 
of N. gentlei, is generally characteristic of older leaves of this plant. One might argue, 
in the absence of data, that the very pronounced pubescence of the younger leaves of 
N. gentlei, as shown in figure 2 in Young (op. cit.), is an adaptation to deter insect 
folivores such as larval P. lypera. DeVries is incorrect in his use of such a labile 
characteristic, in this case, for disclaiming the identification of the food plant. 

DeVries is certainly to be applauded for making the well-founded assertion that one 
needs accurate field data on larval food plants, and that the best data, of course, come 
from complete rearing studies. The incomplete rearing of the larvae in this case, as I 


232 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


thought was explained in Young (op. cit.), was due to running out of food plant when the 
material was brought to Milwaukee, and therefore, the larvae died from starvation and 
not from feeding on the wrong plant. But two events, oviposition, and actual feeding 
on N. gentlei over several days without losses in vigor, together satisfy the criteria for 
an accurate food plant record (Ehrlich & Raven, 1965, Evolution 18:586—608). At the 
very same time, a food plant record, substantiated by recognized authorities in the field 
as exemplified by the association of P. lypera with N. gentlei in northeastern Costa 
Rica, that does not necessarily fit man-made dogma on the coevolutionary interactions 
between plants and insects (lest we believe in a T.C.M., “Tropical Coevolutionary 
Messiah’) should not be rejected as an error. Yet I too agree that there is the need to 
weed out suspect food plant records from the butterfly literature (Ehrlich & Raven, 
op. cit.), something which, based on the data again presented here, DeVries has . 
failed to demonstrate for the butterfly in question. 

I do not question that P. lypera is perhaps polyphagous and indeed feeds on Cap- 
paridaceae as do some other pierids. DeVries apparently has reared P. lypera on this 
family at La Selva. Dr. Hartshorn and I discussing this possibility very briefly tenta- 
tively developed the idea that the greater abundance and density of Nectandra gentlei 
in the premontane tropical rain forest zone encompassing the La Tigra site promotes 
this tree as a frequent food plant of P. lypera at this site, while the interaction may 
shift toward other groups (i.e., Capparidaceae) in lower elevational areas (La Selva), 
where N. gentlei is far less abundant (Dr. G. S. Hartshorn, pers. comm., 24 February 
1981). When a preferred larval food plant becomes very scarce locally and the butterfly 
has the physiological capacity to exploit another family of plants, the carrying capacity 
of the environment can be realized, at least in part, by the expression of polyphagous 
feeding locally in which both families of plants are incorporated into the diet. De- 
pending upon the relative abundance of the two or more plant families exploited by 
the folivore, the biologist studying such a system may encounter one type of interaction 
(food plant association) more frequently than another, particularly when repeated sam- 
ples, as done by DeVries (op. cit.), are taken from the same locality. Current dogma 
may induce one to assume most Neotropical pierids are strictly monophagous, but there 
may very well be cases such as Perrhybris in which ecological factors promote poly- 
phagy. 

I thank the editor of this journal for allowing me the opportunity to make this 
response. I thank Dr. Gary S. Hartshorn, Tropical Science Center, San Jose, Costa 
Rica, for identifying the food plant and for the fruitful discussion. 


ALLEN M. YOUNG, Section of Invertebrate Zoology, Milwaukee Public Museum, 
Milwaukee, Wisconsin 53233. 


Journal of the Lepidopterists’ Society 
36(3), 1982, 233-234 


OBSERVATIONS OF LYCAEIDES ARGYROGNOMON NABOKOVI IN 
THE GREAT LAKES REGION (LYCAENIDAE) 


The presence of the northern blue, Lycaeides argyrognomon Bergatraesser, in the 
Great Lakes Region was first cited by the late Louis Greiswich (1953, Lepid. News 7: 
54), based on a long series of specimens collected in Oconto and Marinette counties, 
1-15 July 1952, in northern Wisconsin. The Greiswich specimens were examined by 
Professor V. Nabokov and found to be an undescribed subspecies of L. argyrognomon. 
It wasn't until 1972 that this population was recognized as a new subspecies, nabokovi 
Masters (1972, J. Lepid. Soc. 26:150-154), occurring in Minnesota and Wisconsin. Mas- 
ters (op. cit.) indicated that the new subspecies may also occur in northern Mich- 
igan. In 1979 James Parkinson (pers. comm.) reported the first capture of L. argyrog- 
nomon nabokovi on 15 July, from Dickinson County, Michigan, thus establishing its 
range in the western portion of Michigan’s Upper Peninsula. Parkinson (pers. comm.) 
again collected specimens of the northern blue on 2-4 July 1980, near Iron Mountain 
in Dickinson County. 

The foodplant of nabokovi was unknown to the authors until 1980, when Les Ferge, 
‘Mo’ Nielsen, and Jim Parkinson discovered females ovipositing on the stems of dwarf 
bilberry, Vaccinium caespitosum Michx., on 4 July, in Florence County, Wisconsin. 
Several females were observed at close range ovipositing on the dwarf bilberry in a 
trailside opening in a wooded area of aspen, oak and jack pine. Oviposition took place 
in the late afternoon between 1500-1730 CDT. The immediate habitat consisted of 
miscellaneous low plants, including grasses and sedges (Carex spp.), creeping black- 
berry (Rubus sp.), bracken fern (Pteridium aquilinum L.) with large patches of V. 
caespitosum on a sandy soil (see Fig. 1). The males were more commonly found flying 
along the adjacent trail and ‘puddling’ at moist spots in the trail. Females were collected 
in close proximity to dwarf bilberry, where they were ovipositing or resting on various 
low plants (see Fig. 2). Farther south along the same trail in Marinette County, several 
northern blues of both sexes were also collected nectaring on Alsike clover (Trifolium 
hybridum L..). On 14 July 1979 at mid-day, females were found nectaring on dogbane 
(Apocynum androsaemifolium L.) and yarrow (Achillea millefolium L.) in Florence 
County. 

Ova collected in the field and obtained from a captive female in 1980 remained 
dormant the rest of the summer, indicating that nabokovi eggs overwinter. An egg was 
also found unhatched in the wild on 1 September 1980 by the junior author. 


Fics. 1-2. 1, habitat of Lycaeides argyrognomon nabokovi Masters, Florence Co., 
Wisconsin; 2, Lycaeides argyrognomon nabokovi female ovipositing on Vaccinium 
caespitosum. 


234 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


The authors have since learned from Robert Dana (pers. comm.) that he also 
observed females of L. argyrognomon nabokovi ovipositing on V. caespitosum in 
St. Louis County, Minnesota. His observations occurred on 5 July 1976 when several 
females were ovipositing on dwarf bilberry, or on debris immediately beneath; a few 
additional ova were found by searching the foodplant. Dana also noted that males 
seemed to concentrate their ‘patrolling’ over patches or mats of dwarf bilberry. 

The northern blue, L. argyrognomon scudderi (Edwards) has been reported from 
Ontario, Canada (1979, Toronto Entomol. Assoc. Occ. Pub. 11:48), with a flight period 
and habitat similar to that of nabokovi. Dr. Nick Escott (1979, ibid.) reported scudderi 
also ovipositing on V. caespitosum on 17 July 1977 in northern Ontario. Is is possible 
that the Ontario population may be synonymous with nabokovi? Until a long series of 
each population can be thoroughly examined, we cannot be certain that the two sub- 
species are actually the same. 


ACKNOWLEDGMENTS 


The authors wish to thank Robert Dana and James Parkinson for furnishing us with 
notes of their field observations, and to Parkinson for guiding us in 1980. We also thank 
Dr. John W. Thompson, University of Wisconsin, Botany Department, for verifying the 
identity of V. caespitosum. 


MOGENS C. NIELSEN, Adjunct Curator, Department of Entomology, Michigan State 
University, East Lansing, Michigan 48824; and LESLIE A. FERGE, 2530 Common- 
wealth Ave., Madison, Wisconsin 53711. 


Journal of the Lepidopterists’ Society 
36(3), 1982, 234-235 


OUTBREAK OF ASTEROCAMPA CLYTON (NYMPHALIDAE) IN LOUISIANA 


In the area adjacent to and including East Baton Rouge Parish, Louisiana, the tawny 
emperor butterfly, Asterocampa clyton (Boisduval & Le Conte), has at least 3 broods 
a year, occurring in early June, mid-July to early August and early September. Adult 
specimens have been taken from April through November. This species is usually ex- 
tremely local but can be common in colonies in bottomland forests. In mid-July 1980, 
an unexpected population outbreak occurred in an area observed to extend as far north 
as Hamburg in Avoyelles Parish and as far west as Port Barre in St. Landry Parish. 
Tens to hundreds of butterflies (females were more conspicuous) could be seen daily 
within the city limits of Baton Rouge, where none or few had been previously recorded. 
Large numbers were observed flying across highways and many were killed by auto- 
mobiles. The species was noticed in West Feliciana, West Baton Rouge, Avoyelles, 
Point Coupee, St. Landry and Iberville Parishes surrounding Baton Rouge. It was found 
to be ee common in areas where its food plant, hackberry (Celtis laeviga Willd.) 
occurred. 


VOLUME 36, NUMBER 3 DENS 


Two 10-minute counts were taken on 14 July 1980. The first count was made around 
a mexican ash, Fraxinus berlandieriana A. DC. at 1700 hrs. CST; 105 female and 64 
male A. clyton were counted feeding on sap, which was exuding from wounds caused 
by borers. This count included 3 mating pairs of A. clyton. Four A. celtis (Boisduval 
& Le Conte) and 1 Polygonia interrogationis (F.) individuals were also present. The 
second count at 1900 hrs. CST under and on a fig tree, Ficus carica L., and particularly 
on fallen fruit, revealed 101 female and 60 male A. clyton individuals including 4 pairs 
in copula. One Papilio troilus L., 2 individuals of Euptychia hermes sosybia (F.), 2 of 
P. interrogationis and 2 of Limenitis arthemis astyanax (F.) were also attracted to the 
fermenting fruit. In both counts many other individuals of A. clyton were flying in the 
near vicinity. 

Egg masses were first noticed on 20 July. The number of eggs in 13 masses ranged 
from 27-135 with a mean of 59. An egg parasite, Telenomus sp. (Scelionidae), probably 
T. rileyi Howard, was identified from several of the eggs. The vespid wasp, Polistes 
exclamans (Vier.), and the spined soldier bug, Podisus maculiventris (Say), were ob- 
served preying on larvae on 12 August. The first adults emerged in late August and 
emergence continued into mid-September. They created a second outbreak which was 
not as large as in mid-July. One worn female was captured as late as 22 October. 

A possible clue as to what upset the normal equilibrium of controlling factors for this 
species can be found in the climatic conditions in the Baton Rouge area in June, July 
and August 1980. Excessively high temperatures in June with an average daily tem- 
perature of 82°F, (1.7°F above normal), July 83.7°F, (1.7°F above normal) and August 
82.2°F,, (0.6°F above normal) were experienced. The areas surrounding Baton Rouge 
endured severe drought beginning in late May and lasting through August. Total pre- 
cipitation in Baton Rouge, however, was above normal in June and July, due to heavy 
rains on 19-20 of June and 18 and 21 of July. These conditions have not been duplicated 
in Baton Rouge climatological history (recorded since 1890). Only 1921 parallels 1980 
in high average temperatures for June and July and precipitation in the summer months. 
Average temperatures in 192] were even higher than those in 1980, 83.1°F in July and 
86°F in August. There is no recorded outbreak of A. clyton in 1921, which may be due 
to the extremely high temperatures or failure to report the phenomenon. It appears 
that very high temperatures in connection with rainfall in June 1980 created conditions 
which directly or indirectly fostered the population explosion in mid-July 1980. The 
continued above-normal high temperatures through August and the increased egg crop 
from the mid-July outbreak account for the second smaller population peak in late 
August to mid-September. 

Specimens collected have been deposited in the Louisiana State University, De- 
partment of Entomology Collection (LSUC). I thank Lubomir Masner, Biosystematics 
Research Institute, Ottawa, Ontario for the identification of he scelionid, and Drs. H. 
B. Boudreaux, J. B. Chapin and L. D. Newsom for their help and suggestions with this 
manuscript. 


MICHAEL L. ISRAEL, Department of Entomology, Louisiana Agricultural Experi- 
ment Station, Louisiana State University, Baton Rouge, Louisiana 70803. 


Journal of the Lepidopterists’ Society 
36(3), 1982, 236-237 


NOTES ON SOME SPECIES OF ASTRAPTES 
HUBNER, 1819 (HESPERIIDAE) 


Astraptes fulgerator (Walch) 1775 


Synonymy: mercatus Fabricius, 1793; fulminator Sepp, 1848; misitra Ploetz 1881; 
albifasciatus Rober, 1925; catemacoensis Freeman, 1967, NEW SYNONYMY. 

Type locality: (?) 

Distribution: U.S.A. (Texas), through Mexico, Central America to Argentina in South 
America. 

Remarks: Apparently there are two subspecies involved here, A. fulgerator fulge- 
rator (Walsh) and A. fulgerator azul (Reakirt). Typical fulgerator occurs from southern 
Mexico (Oaxaca and Chiapas) to Argentina. A. fulgerator has the following character- 
istics: wing bases greenish-blue; the hind termen convex; the upper and lower surfaces 
dark brown; the central band on the primaries dislocated at vein 3, and spot 2 not 
conjoined to the cell spot; usually 3 apical spots in the males, 4 in the females; cilia 
in space 1b white; the white basal streak on the lower surface of the secondaries short 
and broad, with a black streak at the extreme base present; and one of the most im- 
portant characteristics is the broad, distinct, white suffusion in space 1b on the lower 
surface of the primaries. A. fulgerator azul (Reakirt), 1866 is the subspecies that occurs 
in Texas and most of Mexico, extending into South America. A. f. azul has the following 
characteristics: the wing bases are blue or violet-blue; the hind termen straight or 
convex; the upper and lower surfaces dark to pale brown; the central band on the 
primaries usually compact, with the spot in space 3 present or absent; usually 4 apical 
spots but may be 3 or 5; cilia in space 1b brown or white; the white basal streak on 
the lower surface of the secondaries may be short and broad or long and narrow, with 
the black streak at the extreme base of the costa present or absent; and there is usually 
no white suffusion in space 1b on the lower surface of the primaries; however, in the 
blend zone of fulgerator and azul in Oaxaca and Chiapas, Mexico and in Central 
America, this suffusion will be present but never as broad and distinct as is found in 
typical fulgerator. Most of the specimens of azul that I have examined from South 
America have this white suffusion fairly well developed. 

When I described A. catemacoensis from specimens collected at Catemaco, Veracruz, 
Mexico (Freeman, 1967, J. Lepid. Soc. 21:115-119), the description was based on in- 
dividuals that were generally larger and darker than normal fulgerator azul from other 
Mexican localities and also by their having the termen of the secondaries straight. At 
the time I had not examined as many hundred specimens of azul from Mexico as I 
have since and can readily detect that azul is extremely variable and catemacoensis 
should have never been named. With this information I place A. catemacoensis as a 
synomym of A. fulgerator azul. 


Astraptes crana Evans, 1952 


Synonymy: escalantei Freeman, 1967, NEW SYNONYMY. 

Type locality: San Geronimo, Guatemala. 

Distribution: Southern Mexico to Panama. 

Remarks: There are apparently five species or subspecies involved in the creteus 
(Cramer) complex; however, I have not actually examined some of them and am basing 
my information on data stated by Evans (1952, A Catalogue of the American Hesperi- 
idae, Part II, Pyrginae, Sec 1, London, 170 pp.). They may actually all turn out to be 
subspecies of creteus, but for the present I am going to consider them to be separate 
species. The five are creteus (Cramer), 1780; siges (Mabille), 1903; crana Evans, 1952; 
crilla Evans, 1952; and cyprus Evans, 1952. In describing escalantei I was misled by 
Evans discussion of crana, in which he states that on the under-surface of the primaries 
of that species the extreme base of the costa is brown, followed by white to mid-wing. 
This applies to the two females that I have from Rio Santa Domingo, Chiapas, and 
Presidio, Veracruz, Mexico; however, it did not apply to the two males used in the 
description of escaiantei from Ocozingo, Chiapas, Mexico, as the costa on the under- 


VOLUME 36, NUMBER 3 DO 


surface of the primaries was brown throughout with no white present. There was no 
trace of green iridescence at the base that Evans indicated might be present in siges; 
thus, indicating it was not that species. The genitalia are somewhat similar to Evans’ 
figure of creteus. Since describing escalantei I have acquired another male specimen 
from E. C. Welling, collected at Musté, Chiapas, Mexico, 31 July 1968, which has the 
same characteristics as the two males from Ocozingo used in the original description. 
S. R. Steinhauser (1975, Bull. Allyn Mus. No. 29:1-34) indicated in his article “An 
Annotated List of the Hesperiidae of El Salvador,” that the females that he had col- 
lected were definitely crana, but the males were somewhat similar to escalantei. With 
the available information it appears as if the females of Evans’ crana have the char- 
acteristics that he indicated, but the males of that species lack the white on the costa 
of the lower surface of the primaries. With the available information present, I place 
escalantei as a synomym of crana. 


HUGH AVERY FREEMAN, 1605 Lewis Drive, Garland, Texas 75041. 


Journal of the Lepidopterists’ Society 
36(3), 1982, 237 


PROLONGED PUPAL DIAPAUSE OF ALYPIA OCTOMACULATA 
(AGARISTIDAE) 


A recent note by C. Brook Worth (1979, J. Lepid. Soc. 33(3):166) concerning pupae 
of Citheronia regalis Fabricius, which overwintered twice, prompts this additional note 
on pupal longevity. During the twenty years I have been rearing various species of 
moths I have found that a small percentage of the pupae of some species will diapause 
for two years. I have experienced this phenomenon in some broods, though not in 
every brood, of Hemileuca maia Drury, Ceratomia amyntor Hubner, Eupackardia 
calleta West., Callosamia promethea Drury, C. angulifera Walker, hybrids of prome- 
thea X angulifera, and even three and four year diapause in Saturnia pyri Denis & 
Schiffermuller (Bryant, 1980, Maryland Entomologist 1(4):8-9). Alypia octomaculata 
Fabricius represents the first instance of an agaristid with a protracted diapause. 

On 30 April 1977, while on a collecting trip to the Green Ridge Mountain area of 
western Maryland with the Maryland Entomological Society, I caught a female Octo- 
maculata ovipositing on Vitis sp. Upon returning home, the moth was placed in a 
plastic bag containing leaves of Parthenocissus quinquefolia (L.), where it deposited 
approximately eighty ova. The larvae were reared to maturity on P. quinquefolia and 
fifty pupae were obtained (Bryant, in litt.). Since the Baltimore population is double 
brooded, I had expected the moths to begin emerging in July. Apparently, however, 
the western Maryland population is univoltine, as no moths issued from the pupae that 
summer nor have any emerged during the mid-season flight period in the ensuing years. 

The pupae were left, in plastic shoe boxes at ambient temperatures, throughout the 
summer, fall, and winter of 1977 and on 20 May 1978 moths began emerging. Only 
nine adults were obtained in 1978. The pupae remained in the plastic boxes for the 
rest of 1978 and on 2 May 1979 activity was noticed in the boxes. Twenty-five adults 
emerged in 1979. On the chance that there might still be a few viable pupae among 
the remaining sixteen unhatched individuals, they were left undisturbed for a third 
year. Moths were again noticed flying in the boxes on 23 May 1980. Four adults were 
obtained during the spring flight period in 1980. Convinced that I had seen the last 
live moths from those old pupae, I decided to clean out the boxes but luckily never 
followed through. To my astonishment, a single living female was discovered in one 
of the boxes on 18 May 1981. The boxes will now be observed regularly until a season 
passes with no new emergences, at which point any remaining pupae will be dug out 
of their pupal chambers and examined. 


ROBERT S. BRYANT, 522 Old Orchard Rd., Baltimore, Maryland 21229. 


Journal of the Lepidopterists’ Society 
36(3), 1982, 238-239 


FALSE HEAD BUTTERFLIES: THE CASE OF 
OXYLIDES FAUNAS DRURY (LYCAENIDAE) 


I was most pleased to read the recent paper by R. K. Robbins (J. Lepid. Soc. 34:194— 
208) on the false heads of the underside patterns in certain species of the Lycaenidae. 
The bibliography of his paper is also useful. Interesting as that author's research un- 
doubtedly is, I found it even more interesting that there is such a dearth of real research 
into such a promising and fascinating issue. I had always assumed that false heads 
were clear-cut and well documented. 

The paper evoked memories of my own experiences in the late 1960's in Nigeria 
with a species called Oxylides faunas Drury, a butterfly whose behavior, quite literally, 
adds a further twist to the story. 

Oxylides faunas is common in the darker habitats of the tropical zone, such as dense 
primary forest and especially dense secondary forest. Neither sex ever ventures out in 
open sunshine. They normally fly where there is dense undergrowth and usually stay 
at a height of about one meter. The flight is weak and bumpy, most uncharacteristic of 
a member of the Theclinae. The underside displays a splendid example of a finely 
adorned false head. The species almost invariably settles on large, flat green leaves; 
so, the question of whether it settles head-up or head-down is immaterial. 

The special twist is that when Oxylides faunas lands it flicks itself around 180° a 
fraction of a second before landing, so that the false head now faces in the direction of 
flight, fluttering convincingly in even the mildest breeze. 

As luck would have it I received Robbins’ paper a few days before leaving for Nigeria 
on a business trip, and I hoped to substantiate my recollections of more than 10 years 
ago. Although limited time was on hand, I managed to observe a total of 61 landings 
made by 1 male at Akure, Ondo State, 1 male at Benin, Bendel State and 5 males and 


Fic. 1. A specimen of Oxylides faunas seated on a leaf in secondary forest (Agege, 
Lagos, 14 xii 1980). 


VOLUME 36, NUMBER 3 239 


ON 


FIG. 2. Position of head in relation to direction of flight in 61 observed landings of 
14 specimens of Oxylides faunas in Nigeria. 


ABS 


7 females at 4 mi. NW of Agege, Lagos State (Nov.—Dec. 1980). The results of these 
observations are shown in Fig. 2. 

Even the relatively short series of observations clearly shows a statistically significant 
tendency towards making a complete turn. Fifty-one landings (83%) involved a twist 
of more than 90 degrees, most of which were close to 180 degrees (62%). Compared 
to a random distribution the first of these figures is significant at the 0.001 level (Chi- 
square = 27.85, one degree of freedom). In some of the cases where no turn was made 
the reason almost certainly was that the specimen had been disturbed by me to the 
point when normal landing behavior was abandoned. There is a curious leftward bias 
in the turning behavior, with 62% turning left and only 38% right. This is statistically 
significant at the 0.01 level (Chi-square = 7.23, one degree of freedom). The species 
observed by Robbins turned only after landing and did not do so as frequently as 
Oxylides faunas, but there was also a clear leftward bias. 

I recalled that specimens often walked backwards after landing and had supposed 
that this was a further reinforcement of the false head effect. My current series did not 
support this view. About half the landings were followed by walking, but it was usually 
forwards, and the purpose appeared to be to get the butterfly a better launch position. 
There would usually be a pause of three or four seconds between alighting and the 
start of any walk. 

Given the places where Oxylides faunas lands, I would expect the main predators 
to be praying mantisses and hunting spiders, against both of which a false head should 
offer good protection. 


TORBEN B. LARSEN, 23 Jackson’s La., London N. 6, England. 


Journal of the Lepidopterists’ Society 
36(3), 1982, 240 


BOOK REVIEW 


BUTTERFLIES OF THE AFRO-TROPICAL REGION (Volume 2 of Butterflies of the World), 
by Bernard D’Abrera. 1980. E. W. Classey Ltd. Price approximately $150. 


There are others who will review this book from the viewpoint of a professional 
entomologist; this review is the opinion of an experienced amateur. While I think this 
book is well worth the long wait and the relatively high price, I find myself somewhat 
disappointed. The publishers have led us to believe that “it will virtually replace .. : 
Seitz’ (my quotes from a recent Classey flyer), but I cannot agree with this description. 
There is much that is missing from this volume and it appears as though considerations 
of cost have cut out much of what could have made it a true “replacement” for Seitz. 
This is unfortunate, as the volume is the first attempt in over fifty years to cover all the 
African taxa and as such could have commanded almost any price. 

The main problem with the book is its incompleteness. It purports to include Car- 
casson ’s new catalogue, but the list that appears is implicit, is not synonymic, and does 
not have a complete bibliography. A comparison of Peters’ checklist (1952) and 
D’Abrera indicates that many changes have been made in classification over the past — 
30 years and that a number of species that are described in Seitz and supported in 
Peters are missing from the present volume. One has only an occasional! clue as to 
where they have been placed, and the D’Abrera index does not list subspecies, making 
the task even more difficult. Although the author claims that there are only a few species 
which are shown, the book has far too many instances of “I have not seen ....” For 
the book to be complete such statements should be followed with “but the description 
is as follows ...,” this is unfortunately not the case. The author declines to include 
keys “as they would be largely redundant,’ but in many cases the differences are not 
obvious nor are the illustrations good enough to warrant this type of omission. A small 
point—the volume covers the “Ethiopian region” of Seitz but adds to it the area now 
known as Yemen in Southem Arabia. This complicates the problem of the checklist by 
adding a number of species (particularly in the Pieridae). 

The illustrations, while impressive, could be better. In the first place the registration 
and the color balance are both slightly off. Compare the illustration of Lampides boe- 
ticus in this volume with that in volume I (Australian Region) or with that in Pen- 
nington. While this is perhaps an extreme case it points up a technical problem that I 
hope the publishers will correct in future volumes in the series. The author needs to 
improve the technique of photographing reflectant wings, the glare on many Acraeids 
is distracting. In addition I wonder if page folds across wings were not avoidable. 

The central point of my comments is that this-volume is not a complete reference 
work in itself. One still needs Seitz, Peters, Stempffer, Pennington and several other 
references to do a proper job. Collectors in East and Southern Africa have many ex- 
cellent works readily available; those whose collections are concentrated in West Africa 
are still forced to guess. I find I cannot use the book to make a positive identification 
of a specimen even when I know the identity from other sources. A companion volume 
to D’Abrera, containing the Carcasson synonymic catalogue, keys where necessary, and 
original descriptions of species not figured in D’Abrera would solve most of the prob- 
lem and in my opinion is urgently needed. 

I realize that the preceding comments are somewhat negative. However, it should 
be stressed that in spite of these problems it is still a magnificant volume. The pho- 
tographs of the incomparable collection of the BMNH are worth many times the price 
of the book and the shots of collecting locales are positively “mouth-watering.” I highly 
recommend this book to all lepidopterists, professional and amateur. Mr. D’Abrera 
deserves highest praise for his efforts to bring us up to date on the highly complex taxa 
of a rapidly changing continent. 


NICHOLAS G. CARTER, 5406 Huntington Pky., Bethesda, Maryland 20014. 


Date of Issue (Vol. 36, No. 3): 14 December 1982 


EDITORIAL STAFF OF THE JOURNAL 
THOMAS D. EICHLIN, Editor 


% Insect Taxonomy Laboratory 
1220 N Street 
Sacramento, California 95814 U.S.A. 


Macpa R. PAppP, Editorial Assistant 


_ Dovue.as C. FERGUSON, Associate Editor THEODORE D. SARGENT, Associate Editor 


NOTICE TO CONTRIBUTORS 


Contributions to the Journal may deal with any aspect of the collection and study of 
Lepidoptera. Contributors should prepare manuscripts according to the following in- 
structions. 

Abstract: A brief abstract should precede the text of all articles. 

Text: Manuscripts should be submitted in triplicate, and must be typewritten, 
entirely double-spaced, employing wide margins, on one side only of white, 8% x 11 
inch paper. Titles should be explicit and descriptive of the article’s content, including 
the family name of the subject, but must be kept as short as possible. The first mention 
of a plant or animal in the text should include the full scientific name, with authors 
of zoological names. Insect measurements should be given in metric units; times 
_ should be given in terms of the 24-hour clock (e.g. 6930, not 9:30 AM). Underline only 
_ where italics are intended. References to footnotes should be numbered consecutively, 
and the footnotes typed on a separate sheet. 

Literature Cited: References in the text of articles should be given as, Sheppard 
(1959) or (Sheppard 1959, 1961la, 1961b) and all must be listed alphabetically under 


the heading LITERATURE CITED, in the following format: 


SHEPPARD, P. M. 1959. Natural selection and heredity. 2nd. ed. Hutchinson, London. 
209 pp. 

196la. Some contributions to population genetics resulting from the study of 
the Lepidoptera. Adv. Genet. 10: 165-216. 


In the case of general notes, references should be given in the text as, Sheppard (1961, 
Ady. Genet. 10: 165-216) or (Sheppard 1961, Sym. R. Entomol. Soc. London 1: 23-30). 

Illustrations: All photographs and drawings should be mounted on stiff, white 
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_ EXPLANATION OF FIGURES, with a separate paragraph devoted to each page of illus- 


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Proofs: The edited manuscript and galley proofs will be mailed to the author for 
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Correspondence: Address all matters relating to the Journal to the editor. Short 
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02026 U.S.A. 


PRINTED BY THE ALLEN PRESS, INC., LAWRENCE, KANSAS 66044 U.S.A. 


CONTENTS 


SOIL- AND PUDDLE-VISITING HABITS OF MOTHS. Peter H. 
Adler 0 co NE 


REDUNDANCY IN PIERID POLYPHENISMS: PUPAL CHILLING IN- 
DUCES VERNAL PHENOTYPE IN PIERIS OCCIDENTALIS (PIERI- 


DAE). | Arthur M. Shapiro 0 ! 


ROOST RECRUITMENT AND RESOURCE UTILIZATION: OBSERVA- 
TIONS ON A HELICONIUS CHARITONIA L. ROOST IN MEXICO 
(NYMPHALIDAE). ‘'D. A. Waller & L. E. Gilbert 


EVOLUTIONARY STUDIES ON CTENUCHID MOTHS OF THE GENUS 
AMATA: 2. TEMPORAL ISOLATION AND NATURAL HYBRIDIZA- 
TION IN SYMPATRIC POPULATIONS OF AMATA PHEGEA AND 
A. RAGAzZzZU. V.Sbordoni, L. Bullini, P. Bianco, R. Cianchi, 
E. De Matthaeis ¢: S:; Forestiero 3 3 


COCOONS OF CALLOSAMIA PROMETHEA (SATURNIIDAE): ADAPTIVE 
SIGNIFICANCE OF DIFFERENCES IN MODE OF ATTACHMENT 
TO THE Host TREE. G.P.Waldbauer & J. G. Sternburg __- 


NOTES ON THE ACOUSTIC SIGNALS OF A NEOTROPICAL SATYRID 
BUTTERFLY. Stephanie Kane 1) 3 


RELATIONSHIPS BETWEEN PUPAL SIZE AND SEX IN GIANT 
SILKWORM MOTHS (SATURNIIDAE). Thomas A. Miller, 
Jerry W. Highfill & William J. Cooper __.. | eee 


CLASSIFICATION AND LIFE HISTORY OF ACSALA ANOMALA (ARCTI- 
IDAE: LITHOSIINAE). J. Donald Lafontaine, John G. - Fran- 
clemont & Douglas C. Ferguson —-\ 4 


GENERAL NOTES 
A bilateral gynandromorph of Erynnis horatius (Hesperiidae). Michael 
L. Israel G& James E. Gilek 20000 VU . 


The case of Perrhybris lypera (Pieridae) and the Lauraceae: host-plant 
record or assumption? Philip James DeVries 222.0... 
Perrhybris lypera (Pieridae) feeding on Lauraceae: a response to DeVries. 
Allen:M: Young )2.20008 000 NN 
Observations of Lycaedes argyrognomon nabokovi in the Great Lakes 
region (Lycaenidae). Mogens C. Nielsen & Leslie A. Ferge ___-______ 


Outbreak of Asterocampa clyton (Nymphalidae) in Louisiana. Michael 
Ev Tsra@el ov ei DC ane 


Notes on some species of Astraptes Hubner, 1819 (Hesperiidae). Hugh 
Avery Freeman. (000s oe Oe 


Prolonged pupal diapause of Alypia octomaculata (Agaristidae). Robert 


Si Bryant oe Oh Tas) Ud A ea ee 


False head butterflies: The case of Oxylides faunas Drury (Lycaenidae). 
Torben B. Larsen 


BOOK REVIEWS 


185 


192 


200 


207 


218 


297 
229 
230 
233 
234 
236 
237 


238 


— a eS ee 
ee 


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; 
} 
\ 


Sh oe eee 


Rolie 36 1982 , Neer 4 


ISSN 0024-0966 


JOURNAL 


of the 


LEPIDOPTERISTS’ SOCIETY 


Published quarterly by THE LEPIDOPTERISTS’ SOCIETY 


Publié par LA SOCIETE DES LEPIDOPTERISTES 
Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN 
Publicado por LA SOCIEDAD DE LOS LEPIDOPTERISTAS 


DEDICATED TO THE MEMORY OF 


Lloyd Milo Martin 
1912-1982 


18 March 1983 


THE LEPIDOPTERISTS’ SOCIETY 


EXECUTIVE COUNCIL 


CHARLES V. COVELL, JR., President LINCOLN P. BROWER, 
CLIFFORD D. FERRIS, Vice President Immediate Past President 
ALBERTO Diaz FRANCES, Vice President JULIAN P. DONAHUE, Secretary 
CLAUDE LEMAIRE, Vice President RONALD LEUSCHNER, Treasurer 


Members at large: 


R. L. LANGSTON K. S. BROWN, JR. F. S. CHEW 
R. M. PYLE T. C. EMMEL G. J. HARJES 
A. M. SHAPIRO E. H. METZLER 


The object of the Lepidopterists’ Society, which was formed in May, 1947 and for- 
mally constituted in December, 1950, is “to promote the science of lepidopterology in 
all its branches, .... to issue a periodical and other publications on Lepidoptera, to facil- 
itate the exchange of specimens and ideas by both the professional worker and the 
amateur in the field; to secure cooperation in all measures” directed towards these aims. 

Membership in the Society is open to all persons interested in the study of Lepi- 
doptera. All members receive the Journal and the News of the Lepidopterists Society. 
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members should send to the Treasurer full dues for the current year, together with their 
full name, address, and special lepidopterological interests. In alternate years a list of 
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numbers in each volume of the Journal, scheduled for Hebinasy: May, August and 
November, and six numbers of the News each year. 


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Back issues of the Journal of the Lepidopterists’ Society, the Commemorative Vol- 
ume, and recent issues of the NEWS are available from the Treasurer. The Journal is 
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Journal of the Lepidopterists’ Society (ISSN 0024-0966) is published quarterly by the 
Lepidopterists’ Society, a non-profit, scientific organization. The known office of publi- 
cation is 1041 New Hampshire St., Lawrence, Kansas 66044. Second class tes paid 
at Lawrence, Kansas, U.S.A. 66044. 


Cover illustration: Mature Larva of Eumorpha fasciata Sulzer (Sphingidae) feeding 
on Ludwigia sp. (Jussiaea) in southern Florida, where this hawk moth is generally found 
throughout the year. Original drawing by Mr. John V. Calhoun, 382 Tradewind Ct., 
Westerville, Ohio 43081, USA. 


: 


JOURNAL OF 
Tue LeEpPIDOPTERISTS’ SOCIETY 
Volume 36 1982 Number 4 


Journal of the Lepidopterists’ Society 
36(4), 1982, 241-255 


HABITAT, DIVERSITY AND IMMIGRATION IN A TROPICAL 
ISLAND FAUNA: THE BUTTERFLIES OF 
LIGNUMVITAE KEY, FLORIDA’ 


DENNIS LESTON 


Agricultural Research and Education Center, University of Florida, 
Homestead, Florida 33031 


DAVID SPENCER SMITH 


Department of Zoology and Hope Entomological Collections, 
Oxford, England 


BARBARA LENCZEWSKI 


Department of Biological Science, Florida State University, 
Tallahassee, Florida 32306 


ABSTRACT. An annotated account is presented of the 22 resident and 15 casual 
butterfly species reported from Lignumvitae Key, a small, protected and relatively 
unspoiled island in the Florida Keys. The fauna, largely Neotropical, is a segment of 
that found on the Florida mainland and is analyzed in relation to that area and also to 
Cuba and Andros. The majority of species occur in open habitats, the forest being 
impoverished. The overall faunal reduction as compared with the mainland is probably 
an area effect. Among the species on the Key, Eunica tatila, Hemiargus thomasi and 
Phyciodes frisia are noted as being in decline elsewhere in South Florida. 


Two facets of island faunas are of major interest: their biogeograph- 
ical affinities and their diversity. Additionally, there is the frequent 
exhibition of colonization patterns and faunal change (Simberloff, 
1978). Since published information on the Florida Lepidoptera covers 
one hundred years, we have found these insects to be of great value 
in studying the above related topics. The present work considers the 
butterfly fauna of Lignumvitae Key. In the chain of small islands sit- 
uated in Florida Bay known as the Keys, Lignumvitae is the least 
disturbed by the pressures of man and nature and so is of outstanding 
biological importance (Wilson & Eisner, 1968). The species compo- 


' Florida Agricultural Experiment Station Journal Series No. 4462. 


242, JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


<,  | gi°w 
cad eh : 
aq : Dade 


Lignumvitae Key ee, 


Bf ee ) 10 202i 30 


miles 


Fic. 1. Map of the southern tip of the Florida peninsula, including the Florida Keys. 
Lignumvitae Key is indicated (arrow). 


sition of this island takes on even greater significance when viewed 
in the context of a more extensive study of the faunal changes of 
southern Florida, the Keys and the Bahamas currently in progress 
(Lenczewski, 1980; Leston et al., in preparation). 

Lignumvitae Key covers about 270 acres and is situated about half 
a mile north of the eastern tip of Lower Matecumbe Key in the middle 
Keys, Monroe County (Fig. 1). The Florida mainland is approximately 
20 miles to the north, the coast of Cuba is 125 miles to the south and 
the island of Andros in the Bahamas is 140 miles east. The geological 
composition of the Key is Key Largo limestone, a fossilized coral rock; 
and Lignumvitae is the highest of the Florida Keys, reaching 16 feet 
above mean sea level. 

There are no meteorological data available for Lignumvitae Key, but, 
although situated at 24°55’ north and therefore outside the geophys- 
ical tropics, its climate in the Koppen system is of tropical rain forest. 


VOLUME 36, NUMBER 4 243 


600 ft \ 


Fic. 2. Map of the vegetation zones of Lignumvitae Key. Note the pier and adjacent 
yard on the eastern shore. Numerous paths of indeterminate age extend from the yard 
area through the hammock; these have not been surveyed and are not included here. 


There is a dry period extending from November through April, the 
rest of the year being wet. The absence of adequate figures, especially 
of mean hours of bright sunshine per day, does not permit greater 
precision in the defining of seasons (Leston & Gibbs, 1971). 

The Key has a long history of human occupation, dating back to 
pre-Columbian times, but more recent farming has left little mark save 
for some surviving sisal and a few overgrown coconut palms. The 


244 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Matheson house, dating from 1919, is the only substantial building 
on the Key except for outhouses and a caretaker’s dwelling: these all 
lie within a somewhat barren mown yard with a few ornamental plants 
and nearby refuse dumps. A short pier extends from the yard. The 
Key became the property of the State of Florida when it was estab- 
lished as a Botanical Preserve in 1971. | 

The greater part of the vegetation is forest (Fig. 2), locally known 
as “hammock, comprising about 60 species of trees and larger shrubs, 
almost entirely of West Indian origin. Popenoe and Avery (1972) sug- 
gested that “the Key is covered by one of the finest tropical forests in 
an essentially virgin condition to be found anywhere in Florida.” 


METHODS 

Butterflies were netted and, except for hesperiids required for iden- 
tification, were usually released after capture. A voucher collection 
has been deposited with the Department of Natural Resources on 
Lignumvitae Key. 

One or more of the authors visited the Key, singly or in various 
combinations and sometimes accompanied by a colleague, on ten 
mornings between 8 March 1979 and 10 July 1980. A record of person- 
hours collecting was kept (Fig. 3), and a regular route was followed 
to maximize the area covered. No particular attention was paid to 
larvae but any seen were recorded. A few records of adults seen or 
collected by Ranger Jeanne Parks are included in the checklist and 
so identified (J.P.). 7 

After completion of our fieldwork, our attention was drawn to two 
earlier, unpublished studies, the results of which have been incor- 
porated. R. E. Silberglied (pers. comm.) scored butterflies during sev- 
eral visits between 1967 and 1971 and recorded 18 species. The ab- 
sence of precise dates has limited us to listing only the four species 
not found in our own collecting. C. V. Covell (pers. comm.) visited 
the Key on 14 May 1973 and 15 May 1974, and all his records are 
cited below. One of the present authors (D.S.S.) was a member of Dr. 
Covell’s party on the former visit. 

For comparison, data are included for mainland Dade and Monroe 
Counties and the rest of the Florida Keys, based upon the findings of 
Leston et al. (in preparation), for Cuba (Bates, 1935; Riley, 1975) and 
for Andros, Bahamas (Clench, 1977). 


RESULTS 
Terminology 
We use migrate in its basic meaning: to pass from one place to 
another. This excludes “flitting,” the short-range movement associ- 


VOLUME 36, NUMBER 4 245 


ated with the search for food. A regular migrant is a species whose 
movements show a consistent seasonal pattern, as opposed to casual 
migrants (=transients). 


Checklist of Species 


Satyridae 
1. Euptychia areolata Smith 


Present before 1972 (R.E.S.). Clearly a rare casual on Lignumvitae Key, the species 
occurs around freshwater marshes and grassland habitats on the mainland and the larger 
Keys. It is absent from Cuba and Andros. 


Danaidae 
2. Danaus plexippus Linnaeus 


Hammock edge and yard (30.III.79) also 28.X.76 (J.P.). An uncommon but regular 
non-breeding seasonal migrant. Probably absent between May and early October. Oc- 
curs in mainland Dade and Monroe Counties, throughout the Keys, in Cuba and, spo- 
radically, in Andros. The sparcity of Asclepias species host plants on the Key perhaps 
limits the opportunity for breeding, which may take place on the adjacent mainland in 
the dry period. 


3. Danaus gilippus Cramer 


Northwest shore, 17.V.79—when disturbed, the butterflies flew back out over Florida 
Bay; transition zone, 8.VI.79, numerous; yard and east shore, 10.VII.80, three or four 
only. Probably a non-breeding seasonal migrant, surprisingly absent August through 
April. Occurs in mainland Dade and Monroe Counties, throughout the Keys, in Cuba 
and in Andros, where it was first noted in 1977. When compared with plexippus this 
is scarcely to be considered a migratory species, but there is some evidence for move- 
ment (Harris, 1972). Brower (1962) showed that gilippus and plexippus can compete 
for larval food, and the apparent non-overlap of the two on Lignumvitae Key may reflect 
this. Asclepias is scarce on the Key; however, other Asclepiadaceae such as Cynan- 
chum scoparium and Sarcotremma clausa are indigenous and may perhaps be utilized. 


Heliconiidae 
4. Heliconius charitonius Linnaeus 


rammockero lie, 616111) 30.111., 20.1V.; 17.V., 8.VI.; 18.VII., 13.X1.1979, 18.1., 
10.VII.1980. Also 15.V.74 (C.V.C.). Visits the yard in search of flowers, as on 30.III.79 
at flowers of Carissa macrocarpa and on 18.VII.79 far more frequent at the yard edge 
than in the hammock. In all, the butterflies were present in the yard on five of ten 
visits. New broods occur at least in March and June, and observations on a ‘dormitory’ 
of this species in Miami (T. Smith & D. S. Smith, unpublished) suggest that at least 
five generations may be completed in a year. 

A resident, the zebra is found in the nearby mainland counties, throughout the Keys, 
and as different subspecies in Cuba and Andros. The larval foodplants are Passiflora 
species (Passifloraceae), one or two of which occur on the Key. 


5. Dryas iulia Fabricius 


Yard, 8.III., 16.11I., 30.11I., 20.1V., 17.V., 8.VI., 18.VII. 1979, 18.1., 10.VII.1980. Also 
15.V.1974 (C.V.C.). On 16.III.1979, individuals were at the northwest shore on flowers 
of Suriana maritima; on 30.1II.79 some were feeding upon flowers of Carissa macro- 
carpa in the yard. A new brood was present in June. 

A resident, the Lignumvitae Key subspecies is the same as on the adjacent mainland 
and throughout the Keys. Other subspecies occur on Cuba and Andros (Clench, 1975). 
The larvae feed upon Passiflora species. 


246 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


6. Agraulis vanillae Linnaeus 


Yard, 8.III., 16.111., 30.11I., 20.IV., 17.V., 8.VI., 18.VII., 13.1X.1979; 18.1., 10.VII.1980; 
also 28.X.1976 (J.P.). Only found in the yard, where it visits flowers including Carissa 
macrocarpa and Nerium oleander. Most abundant in May through July, with new 
broods occurring at least in early May and late July. 

A resident, the larval foodplants being Passiflora species. This insect is found in 
mainland Dade and Monroe Counties and throughout the Keys, a distinct subspecies 
occurring in Cuba and Andros. 


Nymphalidae 
7. Marpesia petreus Cramer 


Hammock, 10.VII.1980, also sight records 11.VI.1979, 3.VII.1980 (J.P.). The single 
sighting in 1979 suggested a casual immigrant, but the numbers seen within the ham- 
mock in 1980, when up to four were visible at a time, indicate that this is a resident. 
The butterflies were commonly seen around Ficus aurea (Moraceae), a native fig and 
known larval foodplant. 

M. petreus occurs in mainland Dade and Monroe Counties and throughout the Keys 
but is absent from Andros and Cuba. 


8. Eunica tatila Herrich-Schaeffer 


Hammock, 8.III., I6.111., 30111, 20.1V., 17.V., 8:VI., 18: VIL T3aaSiGialisaie 
10.VII.1980, also 27.X.1976 (J. P.) and 14.V.1973, 15.V.1974—“more common than in 
1973” (C.V.C.). This butterfly never leaves the hammock nor, apparently, does it fly 
above the canopy. It was abundant in March but scarce in May and June, with a new 
brood in July (1979, 1980) producing a smaller population than the February brood. 
Numbers are relatively low through the Fall. 

The larval foodplants, all of which are trees, probably include Gymnanthes lucida 
(Euphorbiaceae). Currently, this butterfly occurs in the Upper Keys between Elliot 
Key and north Key Largo, and there are recent reports from Upper Matecumbe and 
Plantation Keys. However, the Lignumvitae Key population is undoubtedly the largest 
concentration. Although a single fresh specimen was found in a Miami garden in 1979 
(D.S.S.), no populations have been found by us in a search of the larger surviving Dade 
and Monroe mainland hammocks, and the last Everglades National Park example dates 
from 1973 (Lenczewski, 1980). E. tatila is present in Cuba but probably absent from 
Andros—see discussion by Clench (1977). 


9. Junonia evarete Cramer 


Yard, 16.11I., 30.1II., 20.1V., 18.VII.1979; also in transition zone, 20.1V., 18.VII., 
13.X1.1979 and in the hammock, 30.III.1979. A resident, but absent or scarce in May 
through July. The larval foodplants include Lippia nodiflora and Stachytarpheta ja- 
maicensis (Verbenaceae). 

This butterfly occurs on Andros and, perhaps, as a separate subspecies in Cuba (cf. 
Riley, 1975). It is widespread in the Keys but confined to coastal areas on the Florida 
mainland: migrations complicate the distribution pattern. 


10. Anartia jatrophae Johansson 


Seaward edge of the yard, 10.VII.1980. Three specimens were seen within five min- 
utes, but absence of this distinctive species from previous samplings or inspections 
and absence of the larval foodplant, Bacopa monnieri (Scrophulariaceae) indicate that 
this is a non-established casual immigrant. It occurs in mainland Dade and Monroe 
Counties, the Keys, Cuba and Andros. 


11. Siproeta stelenes Linnaeus 


Yard, 30.1II.1979, one, at flowers of Carissa macrocarpa; also one seen 18.VI.1979 
(J.P.). A non-established casual immigrant known from Cuba and, for the past 15 or so 


VOLUME 36, NUMBER 4 247 


years, established in the Keys and mainland south Florida but apparently absent from 
Andros. The larval foodplants are Blechum species and perhaps other Acanthaceae. 


12. Phyciodes frisia Poey 


Yard, §.1)., 16.111., 30.1L1., 20.1V., 18.VII., 13.X1.1979, 18.1., 10.VII.1980; also ham- 
mock, 30.III.1979. Also 14.V.1973 (C.V.C.). Numbers were high in March, reduced in 
April and again up (the result of a new brood) in June. Through the rest of the year, 
the biology of the Cuban crescent was unclear. Adults visited flowers of Carissa in the 
yard in late March and flowers of Pluchea purpurescens in the transition zone in July 
1980. A resident, the larval foodplants include Dicliptera assurgens (Acanthaceae), 
which is common on the Key. The species is found in Andros, Cuba and throughout 
the Keys but on the mainland of Florida is now confined to the Flamingo area in the 
Everglades National Park (Lenczewski, 1980). 


13. Phyciodes phaon Edwards 


Yard, 30.III., 20.1V., 19.VII., 13.X1.1979, 18.1., 10.VII.1980. Absent in May and June 
and scarce in July, this butterfly was most frequent in November and January. It is a 
breeding resident, the larval foodplant being Lippia nodiflora (Verbenaceae). Unlike 
frisia this species never strays from the yard. 

Phyciodes phaon is widespread in southern Florida and the Keys, rare in Cuba and 
absent from Andros. 


Lycaenidae 
14. Strymon melinus Hubner 


Yard, 8.III., 16.111., 30.1II., 17.V., 8.VI.1979. This species is a breeding resident with 
two, perhaps three, broods per year. It is highly polyphagous and its larval foodplant 
has not been detected on the Key, but this plant is undoubtedly one of the common 
yard forbs. The butterfly is common in the Keys and on the mainland, absent from 
Andros and Cuba. 


15. Strymon columella Fabricius 


Transition zone, 13.X1I.1979, 10.VII.1980. On the latter date the butterfly was visiting 
flowers of Pluchea purpurescens. Also 14.V.1973 (C.V.C.). The larval foodplant is prob- 
ably one of the Malvaceae, of which several occur as yard forbs on the Key. The species 
occurs on the mainland and through the Keys, also in Andros and Cuba, but there is 
some uncertainty concerning the identity of the involved subspecies (Clench, 1963). 
On Lignumvitae Key, the paucity of records suggests that columella is a casual im- 
migrant, though the presence of potential foodplants indicates that breeding popula- 
tions may be sporadically established. 


16. Strymon martialis Herrich-Schaeffer 


Yard, 30.11]. and 13.X1.1979. Probably a casual migrant, although the foodplant Tre- 
ma micrantha (Ulmaceae) is said to occur on the Key. Widespread, but never abundant 
in mainland Dade and Monroe Counties and in the Keys; it is also present in Cuba 
and Andros. 


17. Brephidium pseudofea Morrison 


Ronson ezones 16-10. SOUL. 201V., 17.Vi. 8:VE., 18.VIL., 13.X1-1979, 18.1., 
10.VII.1980; also on shore of the yard throughout. Other records: 14.V.1973, 15.V.1974 
(C.V.C.). The butterfly was abundant in June and July, common at other times. It is a 
resident breeding species, the larvae feeding on Salicornia species (Chenopodiaceae). 
Adults seldom move more than a few yards from the larval foodplant, and those noted 
at flowers of Pluchea purpurescens (10.VII.1980) were no exception. 

Brephidium pseudofea is present in saltmarshes along the coast of Dade and Monroe 
Counties and in the Keys, but is absent from Cuba and Andros. 


248 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


18. Leptotes cassius Cramer 


Yard, 8.III., 16.111., 30.111., 18.VII., 13.X1.,1979, 18.1., 10.VII.1980; also in transition 
zone 13.XI.1979, and 14.V.1973 (C.V.C.). This butterfly is seemingly absent April through 
June. It is a resident species, the larvae feeding on flowers of a range of herbs, all 
papilionaceous. It is common in mainland Dade and Monroe Counties, throughout the 
Keys, on Andros and in Cuba. 


19. Hemiargus thomasi Clench 


Yard, 8.II1.1979; transition zone and yard, 16.III.1979; hammock, transition zone and 
yard, common 30.11.1979 and in the same areas 20.IV. and 17.V.1979; transition zone 
and yard, 8.VI. and 18.VII.1979; transition zone, 13.X1.1979; yard, 18.11.1980; hammock 
and transition zone, 10.VII.1980. 

A resident species, the larvae feed upon the green seeds within the inflated calyx of 
Cardiospermum halicacabum (Sapindaceae); this is the same foodplant previously not- 
ed by us on Key Largo (Lenczewski, 1980). On Lignumvitae Key, thomasi is essentially 
a species of the hammock canopy and the transition zone edge of the hammock, at 
times flying further afield in search of flowers. 

The Miami blue is now probably extinct in mainland Dade and Monroe Counties, 
but it occurs throughout the Keys from Key Biscayne and Elliot Key southwards and 
on Andros but is apparently absent from Cuba (but see discussion in Riley, 1975). 


20. Hemiargus ceraunus Fabricius 


Yard, 30.III., 13.X1.1979. The paucity of records suggest that this is a casual immi- 
grant, but as the larval foodplants include a range of weedy Caesalpinaceae and Pa- 
pilionaceae (Abrus, Cassia, Crotalaria and Phaseolus species), establishment is pos- 
sible. The species is common on the mainland and occurs throughout the Keys, also 
on Andros and Cuba (but compare the views of Riley, 1975 and Clench, 1977 as to the 
identity of this species and H. hanno Stoll). 


Papilionidae 
21. Papilio cresphontes Cramer 


Yard, 16.III., 30.111.1979, some at flowers of Carissa macrocarpa; hammock, 17.V., 
8.VI.1979; yard and hammock, 13.XI.1979. Also 22.X.1976 (J.P.); 14.V.1973, 15.V.1974 
(C.V.C.). Larvae have been found on Zanthoxylum fagara and Amyris elemifera (Ru- 
taceae). The captures suggest that there are three broods of this resident. The species 
occurs throughout the Florida mainland and in the Keys but is absent from Andros and 
rare in Cuba. 


Pieridae 
22. Phoebis sennae Linnaeus 


Transition zone and yard, 8.III., 30.1II., 8.VI.1979; yard only, 16.111.1979. A regular 
migrant, perhaps temporarily established. The larval foodplants are usually weedy 
herbaceous Papilionaceae or Caesalpiniaceae. The butterfly is common in mainland 
Dade and Monroe Counties and throughout the Keys, in Cuba and Andros. 


23. Phoebis agarithe Boisduval 


Yard, 8.111.1979; transition zone and yard, 16.111. and 30.111.1979, some at flowers of 
Carissa macrocarpa; 20.1V., 17.V., 8.VI., 13.X1.1979, 18.1., 10.VII.1980. Also 14.V.1973, 
15.V.1974 (C.V.C.). Numbers remain fairly constant throughout the year except for a 
dip in July. It is a resident butterfly, the larvae feeding upon Pithecellobium (Mimo- 
saceae), of which two species occur on Lignumvitae Key. Common in mainland south 
Florida and the Keys, the insect is present also in Cuba and Andros. 


24. Aphrissa statira Cramer 


Present before 1972 (R.E.S.) and regarded by us as a rare casual, the ‘migrant sulfur’ 
(Riley, 1975) is recorded from the Keys, the Florida mainland and Cuba but not from 


VOLUME 36, NUMBER 4 249 


Andros. It is very restricted in southern Florida but certainly breeds locally, its numbers 
possibly augmented by immigration. We follow Riley's generic placement. 


25. Eurema lisa Boisduval and Leconte 


Yard, 13.X1.1979, 10.VII.1980. Only one specimen was found in November; the in- 
sect was apparently absent in January and yet numerous in July 1980. We consider this 
species to have been a casual originally, but now established. The larval foodplants 
comprise a range of Papilionaceae and Caesalpiniaceae. Common throughout southern 
Florida and found through the Keys, the butterfly occurs also in Cuba and Andros, but 
there is some confusion concerning the range of the various subspecies. 


26. Eurema daira Godart | 
Yard, 16.II1., 30.1I1., 20.1V., 17.V., 18.VII., 13.X1.1979, 18.1., 10.VII.1980, also 28.X.1976 


(J.P.). The species was most common in each July. A resident and never found away 
from the weed-riddled grass areas of the yard, the larvae feed on a range of low her- 
baceous Papilionaceae. E. daira is common throughout mainland Florida, the Keys, 
Cuba and Andros. Elsewhere (Smith et al., 1982), we have re-examined the problem 
of seasonal and sexual variation in this butterfly, and have included Lignumvitae Key 
material in the analysis. As in Dade County populations surveyed, palmira-like indi- 
viduals occur on Lignumvitae Key. 


27. Nathalis iole Boisduval 


Before 1972 (R.E.S.); 15.V.1974 (C.V.C.). This species was sought for in our visits 
without result. We regard it, therefore, as a casual species on Lignumvitae Key, perhaps 
becoming temporarily established. It is widespread and often abundant on the main- 
land and in other Keys, Cuba and Andros. 


28. Ascia monuste Linnaeus 


Yard and transition zone, 8.III., 16.III., 30.1II., 20.1V., 17.V., 8.VI., 18.VII., 13.X1.1979, 
18.1., 10.VII.1980, including individuals visiting flowers of Pluchea purpurascens. The 
grey form was present in the June sample. The species is apparently most numerous 
in January through March. While a resident, the numbers may well be enhanced by 
regular immigration. Larval foodplants include a range of herbaceous Capparidaceae 


and Cruciferae. The species is common in south Florida, the Florida Keys, Cuba and 
Andros. 


29. Appias drusilla Cramer 


Hammock, 20.IV., 17.V., 18.VII., 13.X1.1979, 10.VII.1980; also at the hammock-yard 
edge, 17.V. and 18.VII.1979. Also 15.V.1974 (C.V.C.). The butterflies were noticeably 
absent in January and March, most numerous in June and July, with a new brood 
present in June. A resident, the larval foodplants include Drypetes (Euphorbiaceae) 
and Capparis (Capparidaceae) species. This pierid is present in the hammocks of main- 
land Dade and Monroe Counties and in suitable habitats throughout the Keys, in Cuba 
and, rarely, in Andros. 


Hesperiidae 
30. Panoquina panoquinoides Skinner 

Shore edge of yard, 18.VII., 13.X1.1979, at flowers of Conocarpus erecta; 18.1.1980. 
Resident, but numbers always low and confined to the narrow coastal edge of the yard. 
The larval foodplant is Sporobolus virginicus (Graminae). In addition to buttonwood, 
the adults utilize the flowers of Sesuvium portulacastrum. The species is found in 
similar habitats on the mainland, throughout the Keys, on Andros but not currently 
occurring in Cuba, according to Riley (1975). 


31. Asbolis capucinus Lucas 


Yard, 8.III., 30.III., 20.1V., 17.V., 18.VII.1979. A resident, the larvae feed upon the 
introduced Cocos nucifera (Palmae). Originally Cuban, capucinus has been estab- 


250 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


lished on the Florida mainland and through the Keys since about 1950. It is not re- 
corded to date from Andros. 


32. Erynnis zarucco Lucas 


Before 1972 (R.E.S.). This skipper is widespread in the lower Keys and present on 
the mainland and in Cuba, but not Andros. Clearly a casual on Lignumvitae: foodplants 
include a few Papilionaceae. 


33. Urbanus proteus Linnaeus 


Yard and hammock, 16.11.1979; yard and transition zone, 13.X1.1979, 18.1.1980, also 
7.XI. and 12.XI.1976 (J.P.). A regular immigrant, temporarily established and breeding 
in the dry period, November through April. The larval foodplants include a wide range 
of herbaceous Papilionaceae. This species is common in mainland Dade and Monroe 


Counties, throughout the Keys, and is represented by a distinct subspecies in Cuba 
and Andros. 


34. Urbanus dorantes Stoll 


15.V.1974 (C.V.C.). Another casual species, recorded only once from Lignumvitae, 
this skipper occurs in Cuba and Andros and, in the past decade or so, has become a 
common butterfly on the Florida mainland and in the larger Keys. It is markedly sea- 
sonal, and its foodplants include many weedy Papilionaceae. 


35. Polygonus leo Gmelin 


Yard, 16.II1., 30.111.1979, at flowers of Gliricidia and Carissa; 20.IV., 17.V.1979, 18.1., 
10.VII.1980; in hammock, 30.III., 20.I1V., 13.X1.1979, 18.1.1980; also in transition zone, 
13.XI1.1979. Also taken 15.V.1974 (C.V.C.). This hesperiid, a resident, is most common 
November through March. The larvae feed upon Piscidia piscipula (Papilionaceae). 
The distribution includes mainland Dade and Monroe Counties, the Keys, Cuba, but 
apparently not Andros. 


36. Epargyreus zestos Geyer 


Hammock, 10.VII.1980, a single worn individual; also 15.V.1974 (C.V.C.). A casual 
immigrant, now probably extinct on mainland Florida but unpredictably present through 
the island chain from Key Largo to Key West; found on Andros but absent from Cuba. 
The larval foodplants are probably legumes (Riley, 1975). 


37. Phocides pigmalion Cramer 


Yard, 8.II1., 16.111.1979, at flowers of Gliciridia. 30.111.1979 numerous, some at flow- 
ers of Carissa, 20.1V., 18.VII. (new brood), 13.X1.1979, 18.1., 10.VII.1980; also in tran- 
sition zone, 20.1V.1979 and 28.X.1976 (J.P.). A resident, the larvae feed on the leaves 
of Rhizophora mangle (Rhizophoraceae). This skipper is widespread in the belt of red 
mangrove of Dade and Monroe Counties, in the Florida Keys and, as distinct subspe- 
cies in Andros and Cuba. 


DISCUSSION AND ANALYSIS 


The butterfly species found on Lignumvitae Key may be divided 
according to their status as follows: 


Immigrant, casual, non-breeding iil 
Immigrant, regular, breeding 1 
Immigrant, regular, non-breeding ©) 
Immigrant, total 15 
Resident, breeding, total Ue) 
Species total 37 


The species, immigrant and resident, also segregated by habitat: 


VOLUME 36, NUMBER 4 Ob 


Immigrant Resident Total 
Forest: hammock 1 6 Uf 
mangrove 0 1 1 
total AS) 
Open: — yard/scrub 9 13 22, 
shore/saltmarsh 0 2 2) 
total 24 
Total (all habitats) 3) 


Species noted by previous observers (E. areolata, A. statira, N. iole, 
E. zarucco and U. dorantes) for which locality data is not available 
are not included in the above analysis. Although not statistically sig- 
nificant, it is noteworthy that all but one of the hammock species are 
resident. There is a marked preponderance of species of open habi- 
tats, though these comprise less than 20% of the Key’s area: the 50% 
probability test indicates a significant departure from a random dis- 
tribution hypothesis—0.05 > P > 0.01. Based on mainland observa- 
tions of habitat preference, none of the five species mentioned above 
is a forest dweller, suggesting that the significance of non-random 
distribution based on our survey is underestimated. 

The Lignumvitae Key species, resident or immigrant, occur in 
neighboring areas in the following numbers: 


Florida Mainland 
Lignumvitae Keys So. Florida Cuba Andros 
Numbers a7 ove 34 30 5) 
Percent 100 100 91.9 81.1 67.6 


The butterfly fauna of Lignumvitae Key contains no unique species, 
all being found elsewhere in the Florida Keys, but beyond this island 
chain, there is a reduction with distance of species held in common. 
The difference from the nearby Florida mainland (Dade and Monroe 
Counties) is the result solely of the recent extinctions in the Miami/ 
Homestead area: E. tatila, H. thomasi and E. zestos have been lost 
within the past ten to twenty years (Lenczewski, 1980; Leston et al., 
in prep.). Phyciodes frisia too, once widespread in Dade and Monroe 
Counties, is now confined to the area around Flamingo, Everglades 
National Park and is another species of diminishing range. 

It is assumed that Lignumvitae Key depends for its faunal diversity 
upon the pool of species provided by the adjacent Florida mainland 
and the chain of islands of which it is a part. Leston et al. (in prep.) 
give a checklist of the present day butterflies of Dade and Monroe 
Counties, which forms the basis of the following comparison; the 
higher groupings, for convenience follow Klots (1951). Where a species 
on the mainland comprises both breeding residents and migrants (e.g. 
Danaus plexippus and Ascia monuste) it is scored in the former cat- 
egory. 


252 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Dade and Monroe Counties Lignumvitae Key 
Family Resident Migrant Total Resident Migrant Total 
Satyridae 2 0 2 0 i i 
Danaidae 3 IL 4 0 2 2) 
Heliconiidae 3 0 3 3 0 3 
Nymphalidae 12 13 25 5 2 i 
Libytheidae 0 1 IL 0) 0 0 
Riodinidae 1 0 1 0 0 0 
Lycaenidae 16 0 16 4 3 a 
Papilionidae 6 6 12 ] 0 1 
Pieridae 13 7 20 5 3 8 
Hesperiidae 36 fi 43 4 4 8 
Totals 92 35 D7 22 1,53 on 


Spearman s rank correlation test shows there is a significant positive 
correlation between families of the mainland plus Keys fauna, on the 
one hand, and that of Lignumvitae on the other: >} d? + t = 20.5, n = 
10, P < 0.01. The assumption made above is therefore confirmed. 

The proportion of migratory to resident species is higher in the 
Lignumvitae list than in the species pool: 


Resident Migratory Total 
Dade and Monroe 92 (72.4%) 35 (27.6%) 127 
Lignumvitae Key 22 (59.5%) 15 (40.4%) oe 
L.K. fauna as : 
percent of pool 23.9% 49.9% 29.1% 


Overall, the butterflies of Lignumvitae Key represent a little over a 
quarter (29.1%) of the species pool in Dade and Monroe Counties. 

The butterfly fauna of the Key has 31 species whose affinities are 
Neotropical, with only 6 (S. melinus, P. cresphontes, E. areolata, E. 
lisa, N. iole, E. zarucco) with Nearctic affinities. 

The relationship of collecting hours to the cumulative number of 
species noted is shown in Fig. 3. It suggests that additional species, 
probably non-resident, remain to be recorded for Lignumvitae Key. 


DISCUSSION 


The butterflies of Lignumvitae Key are overwhelmingly of Neo- 
tropical origin, but the absence of Cuban or Bahamian species not 
found on the Florida mainland suggests that the island has probably 
been colonized, not directly from Cuba and/or the Bahamas, but from 
the neighboring Florida Keys and peninsular Dade and Monroe 
Counties. This is scarcely surprising in view of the relative distance 
involved and applies whether we consider resident or immigrant 
species separately or in combination. 


VOLUME 36, NUMBER 4 253 


40 
30 


20 


cumulative spp. 


10 
10 30 50 70 


hrs collecting 


Fic. 3. Illustrating the relationship between person/hours collecting and cumula- 
tive number of species recorded (logarithmic). 


We meet a phenomenon on the Key of species occurring as tran- 
sients (casual immigrants) which, in the much larger area of the south 
Florida mainland and even on some of the larger Keys, are breeding 
residents: examples include Danaus gilippus, Anartia jatrophae and 
Siproeta stelenes. MacArthur and Wilson (1967) hypothesized that 
this might happen, though with little factual data in support. How- 
ever, whether the absence of species common elsewhere in the vi- 
cinity or whether some species are transient instead of permanent can 
be construed as the result of direct area effect is not at first evident 
from our data. Simberloff (1978) avers: “... area affects species num- 
ber independently of habitat diversity.”” The absence from Lignum- 
vitae Key of such important components of the nearby land areas as 
pinelands, oak hammocks, Everglades prairies, freshwater marshes 
and agricultural plots, each of which supports a characteristic butterfly 
fauna, may directly limit species diversity, though the addition of only 
one or two of these biotypes can be envisioned in an area as small as 
270 acres. 

If we ignore exact species composition and work at a higher taxo- 
nomic level, as expressed by the list of families and their relative 
numbers of included species, the significant positive correlation not- 
ed indicates that the Lignumvitae Key butterfly fauna may be viewed 
as a reduced but undistorted simulacrum of the pool community. In 
other words, direct area effects rather than habitat simplification, may 
suffice to explain the differences between the faunas of the pool and 
of the Key. 

Both from a study of succession (Southwood et al., 1979) and from 
a direct comparison of the phytophagous insect fauna of tropical forest 


254 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


with that of non-forest (Leston, 1980), the wooded climactic areas of 
Lignumvitae Key would be expected to support more species than 
the open places. In fact, the reverse is the case: we find the forest of 
this Key, like the hammocks of mainland south Florida, to carry an 
impoverished butterfly fauna. 

The introduction of Papilio aristodemus ponceanus (Schaus’ swal- 
lowtail) to the Key has been contemplated (Covell, 1976). The prin- 
cipal hostplant of this species Amyris elemifera (torchwood) is pres- 
ent on this Key, though sparsely, but the butterfly has not been reliably 
reported from Lignumvitae or, since May 1945, from nearby Lower 
Matecumbe Key (Henderson, 1946). P. a. ponceanus is established in 
a threatened area on north Key Largo and is under effective protection 
on Elliot Key and other islands in the Biscayne National Monument. 
Covell (1976) has stressed the fluctuation in abundance of this species 
from year to year: we cannot predict its long-term future in the area, 
but should a further protected habitat be needed to aid its conserva- 
tion, Lignumvitae Key remains the obvious choice. In addition, it 
should be noted that Lignumvitae Key possesses one of the best sur- 
viving U.S. colonies of Eunica tatila, a species that declines as de- 
struction of hardwood forest progresses. Other species well estab- 
lished on the Key, but with diminishing range elsewhere in the area 
and notably on the south Florida mainland, include Phyciodes frisia 
and Hemiargus thomasi. 


ACKNOWLEDGMENTS 


We thank Major James A. Stevenson (Division of Recreation and Parks, Florida) for 
permission to carry our research on Lignumvitae Key, the staff at Long Key State 
Recreation Area and Lignumvitae Key State Botanical Site for supplying transportation 
and for their generous support in many ways. Particularly, we thank Ranger Jeanne M. 
Parks for advice, help and the use of her personal records. We also thank Frank Rivera 
(Dade County Parks Service) for his assistance in the field, Dr. Richard M. Baronowski 
(University of Florida, Homestead) for first introducing one of us (D.L.) to the problems 
of the Key, and for much useful discussion. We thank Dr. Daniel S. Simberloff and Dr. 
Donald R. Strong (Florida State University) for their constructive criticism of the manu- 
script, and we especially thank Dr. Robert E. Silberglied (Harvard University) and Dr. 
Charles V. Covell (University of Louisville) for providing us with lists of their captures 
and sightings and copies of their surveys. 

We are sad to record that Dr. Dennis Leston, who initiated the survey described in 
this report, died in October 1981. An appreciation of his work has been provided by 
T. R. E. Southwood, 1982. Antenna 6:173-174. 


LITERATURE CITED 


BATES, M. 1935. The butterflies of Cuba. Bull. Mus. Comp. Zool. 78(2):63-258 

BROWER, L. P. 1962. Evidence for interspecific competition in natural populations 
of the monarch and queen butterflies, Danaus plexippus and D. gilippus berenice 
in south central Florida. Ecology 43:549-552. 


VOLUME 36, NUMBER 4 255 


CLENCH, H. K. 1963. A synopsis of the West Indian Lycaenidae, with remarks on 
their zoogeography. J. Res. Lepid. 2(4):247-270. 
1975. Systematic notes on Dryas iulia (Heliconiidae). J. Lepid. Soc. 29:230- 


230) 

. 1977. A list of the butterflies of Andros, Bahamas. Ann. Carnegie Mus. 46(12): 
173-194. 

COVELL, C. V. 1976. The Schaus Swallowtail: A threatened subspecies? Ins. World 
Digest 3:21-26. 

HarRRIS, L. 1972. Butterflies of Georgia. Norman: Univ. Oklahoma Press. 

HENDERSON, W. F. 1946. Papilio aristodemus ponceanus Schaus (Lepidoptera: Pa- 
pilionidae) notes. Entomol. News 57:100-101. 

Kiots, A. B. 1951. A Field Guide to the Butterflies of North America, East of the 
Great Plains. Boston: Houghton Mifflin. 

LENCZESWKI, B. 1980. Butterflies of Everglades National Park. Homestead: South 
Florida Research Center. 

LESTON, D. 1980. The natural history of some West African insects. 14. Entomological 
changes accompanying degradation of the forest. Entomol. mon. Mag. 155:105- 
116. 

& D. G. Gipsps. 1971. Phenology of cocoa and some associated insects in 
Ghana. Proc. 3rd Int. Cocoa Res. Conf. Accra 1969:197-204. 

MACARTHUR, R. H. & E. O. WimSON. 1967. The Theory of Island Biogeography. 
Princeton: Princeton Univ. Press. 

POPENOE, J. & G. AVERY. 1972. Survival of ornamental plants on Lignumvitae Key. 
Proc. Florida State Hort. Soc. 84:368-370. 

RitEy, N. D. 1975. A Field Guide to the Butterflies of the West Indies. London: 
Collins. 

SIMBERLOFF, D. S. 1978. Colonisation of islands by insects: Immigration, extinction 
and diversity. Symp. Royal Entomol. Soc. London 9:139-153. 

SMITH, D. S., D. LESTON & B. LENCZEWSKI. 1982. Variation in Eurema daira (Lep- 
idoptera: Pieridae) and the status of palmira in Southem Florida. Bull. Allyn Mus. 
No. 70:1-8. 

SOUTHWOOD, T. R. E., V. K. BROWN & P. M. READER. 1979. The relationships of 
plant and insect diversities in succession. Biol. J. Linn. Soc. London 12:327-348. 

THE TIMES. 1968. The Times Atlas of the World. London: Times Newspapers. 

WILSON, E. O. & T. EISNER. 1968. Lignumvitae: Relict island. Natural History 77(8): 
52-57. 


Journal of the Lepidopterists’ Society 
36(4), 1982, 256-263 


FIELD OBSERVATIONS OF FOODPLANT OVERLAP AMONG 
SYMPATRIC CATOCALA FEEDING ON JUGLANDACEAE 


DALE F. SCHWEITZER 


Curatorial Affiliate, Entomology Division, Peabody Museum of Natural History, 
Yale University, New Haven, Connecticut 06511 


ABSTRACT. Field collections or observations of eggs, larvae and ovipositing fe- 
males confirm that Catocala epione Drury, habilis Grote, judith Strecker, obscura 
Strecker, residua Grote, retecta Grote, and palaeogama Guenée feed on shagbark 
hickory, Carya (section Eucarya) ovata in Connecticut. C. obscura and probably C. 
residua prefer this species over other Eucarya while C. palaeogama seems to oviposit 
randomly on species of this section. No species of Catocala appears to prefer a species 
of Eucarya other than C. (E.) ovata. None of the Eucarya feeders were encountered 
on section Apocarya or Juglans. Since C. ovata accounts for well over half of all Jug- 
landaceae in Connecticut, it would be the de facto primary foodplant of any Catocala 
species that oviposits randomly on Eucarya. The hypothesis that Catocala are exten- 
sively partitioning the available Eucarya by species is untenable. Individual trees are 
frequently used by several Catocala species, and there is little evidence of partitioning 
at that level except that C. epione appears to feed largely on small plants. 

Catocala subnata Grote oviposits on Carya (section Apocarya) cordiformis and Ju- 
glans cinerea. Catocala neogama (J. E. Smith) and C. piatrix Grote probably feed 
largely or entirely on Juglans in Connecticut. Few comparable assemblages of oligoph- 
agous or monophagous congeneric Lepidoptera utilizing a single foodplant occur in 
the North Temperate Zone. 


The moth genus Catocala is well known for its extreme sympatric 
diversity (see Sargent, 1976). In southern New England, three well 
sampled locations have produced from 35 to 40 species! each (Sargent, 
1976), and I have taken 39 species in seven years within a kilometer 
of my home in Hamden, Connecticut, almost entirely at sugar bait. 
Most of the species are taken every year, and many are taken in large 
numbers. I estimate that 52 species of Catocala have been taken in 
Connecticut since about 1890. Forty-six of those are present now on 
a regular basis. At least one that is now common (C. judith Strecker) 
is absent from old area collections, while C. robinsonii Grote was 
established between about 1898 and 1964, but no longer occurs. C. 
pretiosa Lintner has also disappeared. 

In addition to sympatry, Catocala have been suspected of extensive 
foodplant overlap. For example, fourteen species of current Connect- 
icut Catocala are well known to use Juglandaceae as larval foodplants. 
Four additional Juglandaceae feeders have been taken in the state. 
However, the foodplant compilations of Sargent (1976) are based 
largely on rearings ex ovis, and it is not possible to determine precise 
natural foodplants. The situation with older references is much worse 
as specific plant names were almost never given. Thus, with two 


'C. “amica” includes two species at all three localities. 


VOLUME 36, NUMBER 4 257 


species of Juglans and five of Carya [four Section Eucarya, one Apo- 
carya (Fernald, 1950)] present widely in Connecticut, it seemed pos- 
sible that fourteen Catocala species indeed might be partitioning the 
Juglandaceae by species to some extent. However, one species of 
hickory, Carya ovata, greatly outnumbers all other combined Juglan- 
daceae in most portions of the state (see localities, below). 

With the data presented below, I intend to document the Catocala 
fauna of Carya ovata and to provide some indication of the range of 
foodplant choices of these moths. 


METHODS 


A variety of methods have been used to document foodplant usage 
in the field. During the winter of 1978-1979 bark samples of five to 
ten shags were taken from seven mature Carya ovata and two mature 
C. glabra at Roxbury, Litchfield Co., Connecticut, as well as from two 
to five mature C. ovata each from four localities in New Haven Coun- 
ty, for a total of 18 C. ovata samples, each containing visible Catocala 
eggs. 

Bark samples were sleeved on C. ovata in April 1979, and larvae 
were changed to new sleeves as needed in June. A limited supply of 
sleevable branches necessitated pooling of some samples. A more 
serious complication was the excessive number of larvae produced. 
The two most rapidly growing species, C. palaeogama and C. residua 
severely depleted food supplies, and this may well have led to high 
mortality of other species. Table 1 gives the results of this method. 

A second method was searching for larvae. On 30 June 1979 I 


TABLE 1. Catocala reared from eggs collected during the winter of 1978-1979 in 
Litchfield (Roxbury) and New Haven (all others) Cos., Connecticut. Identifications 
were mostly from adults, but some were from larvae which were subsequently pre- 
served. All eggs were sleeved on Carya ovata. 


Catocala produced 


Locality and tree palaeogama residua retecta obscura 


Carya ovata 


Roxbury, tree B 6 3 i 0 
‘G PN tS 0 0 
D ? 2, 0 0 
E 13 6 0 0 
Roxbury, trees A, F, G ao 14 4 2 
Lake Gaillard, 2 trees 20 0 0 0 
New Haven-Hamden, 9 trees ad. 0 2 0) 


Carya glabra 
Roxbury, 2 trees 6 0 0 0 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


258 


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VOLUME 36, NUMBER 4 259 


TABLE 3. Summary of Catocala ovipositions and number of trees (Carya, Juglans) 
checked in New Haven County, Connecticut (during 1979 and 1980). Numbers in 
parentheses refer to same night repeats of previously checked trees (see text). 


Catocala species 


N obser- palaeo- 
Tree species vations neogama subnata gama _ residua obscura judith  retecta 
C. ovata 179 (12) ) 0 42 IS) Sap) 1 3 
C. glabra 85) (3) 0 0) 4 Ik 0 0 0 
C. ovalis 5 0 0 1 0 0 0 0 
C. tomentosa 13 0 0 3 0 0 0 0 
C. cordiformis 36 (19) 0 5 0 0 0 0 0 
J. cinerea 19 (10) il il 0 0 0 0 0 


searched for larvae under shags of mature C. ovata near North Ash- 
ford, Connecticut. No other Carya species was encountered, but four 
Juglans cinerea were checked. By this date most larvae had probably 
pupated. During May 1980 young larvae were collected from foliage 
and branches of small hickories on West Rock, New Haven, Con- 
necticut. Branches were beaten after being inspected visually. Most 
larvae were found by inspection. Larvae were also taken on Juglans 
in various years on the trunk, at the base of large limbs, or in debris 
at the base of the trees. None were found on small branches or leaves. 
Numbers of trees searched and larvae found are given in Table 2. 

A third method was searching for ovipositing females at night when 
they are easily found on tree trunks. Initial attempts at this method 
in 1979 were somewhat haphazard, but systematic searches were im- 
plemented that year and in 1980 with exact numbers of trees searched 
being recorded (Table 3). 

Tree trunks were searched after dark with lights, and when possi- 
ble, moths were captured. Only females seen probing bark crevices 
with the tips of their abdomens were recorded as ovipositing. Inter- 
estingly, startled females sometimes returned to such activity almost 
immediately. Very few females were encountered that were not clear- 
ly ovipositing. The most effective procedure was to have one person, 
armed with a net and cyanide jar, search using a headlamp, while a 
second person would operate a more powerful hand-held lantern and 
record data. 

Some trees were checked more than once as indicated in Table 3. 
Such checks were always more than forty-five minutes apart, and moths 
were removed each time; therefore, I treat these as independent ob- 
servations. All Juglandaceae over about 5 cm in diameter that were 
encountered were checked. 


260 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Localities 


All localities at which searches for ovipositing females were con- 
ducted are in New Haven County, Connecticut. The majority of ob- 
servations were on Nelson Rd., Southbury, or on West Rock Ridge, 
New Haven. Other observations were from Brooksvale Park, Hamden 
and various areas in Cheshire and Wallingford. Except on West Rock, 
all trees were mature, almost always with trunk diameters of 25 cm 
or more. Some were greater than 70 cm in diameter. Most of the trees 
were at the edges of extensive mixed mesic forests. 

West Rock Ridge is a hotter and drier habitat than the others with 
thin acid soils overlying trap rock. The hickories there are stunted, 
though numerous. Very few have diameters of 20 cm or greater. The 
canopy is more open than at the other forested sites. This is the only 
one of the localities where C. ovata accounted for less than half of 
all Juglandaceae searched, and the only one where Carya ovalis was 
seen. 

At Brooksvale Park, Hamden the area checked contained 13 C. ova- 
ta, 1 C. glabra, 2 C. tomentosa and 1 J. cinerea. The Southbury area 
contained 36 C. ovata, 9 C. glabra, 6 C. cordiformis and 3 J. cinerea. 
At West Rock 22 C. ovata, 16 C. glabra, 5 C. ovalis, 6 C. tomentosa 
and 1 J. cinerea were checked. The Wallingford area contained 10 C. 
ovata, 1 C. glabra, 1 C. tomentosa, 4 J. cinerea, and the Cheshire 
locality had 8 C. ovata and 1 J. cinerea. Most of these areas were 
checked on more than one occasion, and some trees were sometimes 
overlooked. 

The few larval data from New Jersey are from the vicinity of villages 
in the Pine Barrens region, ca. 200 kilometers southwest of the Con- 
necticut sample areas. 


RESULTS 


Egg and larval samples. These data (Tables 1 and 2) permit several 
conclusions: 1) at least Catocala epione, habilis, obscura, residua, 
retecta and palaeogama utilize Carya ovata as an important foodplant 
in Connecticut; 2) C. neogama is restricted, or nearly so, to Juglans 
in this region; 3) those species regularly utilizing Carya were not 
encountered on Juglans; 4) little can be deduced regarding utilization 
of other Carya spp., but C. palaeogama at least uses species other 
than C. ovata in Connecticut and C. epione does so in New Jersey. 

It was not uncommon for several species to occur together on in- 
dividual trees. C. epione seems to utilize primarily small plants (based 
on observations herein, and those of H. D. Baggett, L. F. Gall, T. D. 
Sargent, all pers. comm.), but the other Carya feeders must often all 
occur together. 


VOLUME 36, NUMBER 4 261 


One unsolved enigma is the absence of C. habilis, a common species, 
from oviposition observations (below) and bark samples. L. F. Gall 
(pers. comm.) has also failed to record it. Perhaps it oviposits very 
late at night or high in the trees. Larvae are common on Carya ovata 
(Table 2; L. F. Gall, pers. comm.). 

Analysis of oviposition data. The data in Tables 2 and 3 convinc- 
ingly demonstrate that C. palaeogama, C. residua, C. obscura and C. 
retecta regularly utilize Carya ovata for oviposition. C. judith is also 
added to the list of C. ovata feeders. 

Chi-square goodness of fit tests were performed with oviposition 
data for those Catocala species for which ten or more observations 
are available. These analyses show that oviposition is non-random 
among all Juglandaceae for C. palaeogama (P < .001), residua (P < 
.O1), and C. obscura (P < .001). C. palaeogama apparently oviposits 
randomly on any Eucarya (P > .20), while C. obscura prefers C. ovata 
over the other (pooled) Eucarya (P < .05). I suspect that C. residua 
also prefers C. ovata over other Eucarya in this region, but current 
data do not demonstrate such a preference (P > .10), probably due to 
the small sample size. The preferences of C. palaeogama (P < .001) 
and C. obscura (P < .05) for Eucarya over Apocarya and Juglans com- 
bined are significant. 

Data for C. subnata are not adequate for analysis, but it is pre- 
sumably limited to Apocarya and possibly Juglans. Juglans cinerea is 
the only previously reported host (Sargent, 1976). In 1981 L. F. Gall 
and I attempted to rear the progeny of three different female C. sub- 
nata. Hatchlings accepted Carya (Apocarya) cordiformis, and adults 
were obtained using this plant. C. (A.) illinoiensis was also readily 
accepted, but the plants available were too small to allow for an at- 
tempt at rearing larvae on this species. In no-choice situations, some 
larvae would accept both species of Juglans, but no larvae matured 
in sleeves on these. When cuttings were used, no larvae completed 
the first instar. My larvae would not accept any species of Eucarya. 
In 1981 I observed two more C. subnata ovipositing on C. cordiformis 
and L. F. Gall also observed several. Aside from C. subnata, no species 
of Connecticut Catocala seems to prefer a species of Carya other than 
C. ovata. 


DISCUSSION 


The above observations document that not less than seven species 
of Catocala utilize Carya ovata in Connecticut. These are C. epione, 
C. habilis, C. judith, C. obscura, C. residua, C. retecta and C. pa- 
laeogama. I have also reared adults of all of these ex ovis on C. ovata. 
Catocala serena Edwards adults rest on this tree in Connecticut and 


262 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


have been observed ovipositing on it in Michigan (Nielsen, 1978). 
Sargent (1976) and myself have reared C. dejecta Strecker on Carya 
ovata, and in some of the moth’s localities other hickories are nearly 
absent. Juglans cinerea is also an acceptable foodplant for at least the 
first three instars (pers. obs.). C. dejecta is very rare in New Haven 
County where oviposition observations were made. Two other Carya 
feeders, C. vidua (J. E. Smith) and C. flebilis Grote, occur in Con- 
necticut at present. Their local foodplants are unknown. Thus, it is 
quite possible that 11 species of Catocala use Carya ovata in Con- 
necticut and, as discussed previously, some of these species prefer 
this plant. However, even random oviposition on any Eucarya indi- 
vidual would result in C. ovata being de facto the principal foodplant 
in most of the state (see localities above). Thus, the hypothesis that 
these Catocala extensively partition the Eucarya by species is unten- 
able. 

There is at present no evidence that they are extensively partition- 
ing C. ovata by individual trees, except for C. epione, although more 
data are desirable. The four trees on which multiple (>2) ovipositions 
were observed always had more than one species present. At South- 
bury on 2 September 1980 one tree had three C. palaeogama, one C. 
judith and one C. obscura ovipositing on it, while an adjacent tree 
had two C. palaeogama, two C. obscura and one C. retecta. Obser- 
vations of C. residua and C. palaeogama ovipositing together have 
been frequent, and at least three species have been obtained from 
some larval collections and bark samples (Tables 1, 2). Doubtless, had 
observations of individual trees been carried out over the entire Ca- 
tocala season at Southbury (the early C. judith was largely missed), 
many trees would have had recorded ovipositions by five or more 
species. 

Although the observations reported here show that several species 
of Catocala larvae frequently occur together on Carya ovata, there 
are insufficient data to determine whether other forms of niche split- 
ting are used. With seven or more species potentially occurring on 
the same tree and with all of these feeding exclusively on foliage and 
most having similar egg hatching dates (manuscript in prep.), such 
niche splitting would have to be rather fine. Sargent (1976) has point- 
ed out that some similar species grow at different rates as larvae, and 
I have similar data (unpublished). Eucarya foliage appears rather syn- 
chronously in early spring, and additional foliation after early May is 
rare. Therefore, late maturing species (e.g. C. habilis, obscura) might 
compete with the early species if food became limiting. Effective 
avoidance of competition for food, if such occurs, would probably 
have to involve utilization of different portions of the tree. As I note 


VOLUME 36, NUMBER 4 263 


elsewhere (Schweitzer, 1982) there is some evidence for differences 
in resting behaviors (see also Sargent, 1976). 

Casual observation suggests that Catocala larvae are held at rela- 
tively low numbers (relative to leaf availability) by some as yet un- 
known factor, probably predators and/or parasitoids. One can consult 
most ecology texts (e.g. Ricklefs, 1973, pp. 510, 522-523) and find that 
the “principle of competitive exclusion” applies only with regard to 
limiting resources. Sargent (1976, 1977) suggests foodplant availabil- 
ity may not be an important factor in Catocala evolution. The data in 
this paper support that contention at least to the extent that compe- 
tition has not forced the Eucarya feeders to specialize on different 
species or to avoid frequent co-occurrence on individual trees. 

Nevertheless, I am not aware of any well documented comparable 
sympatric assemblages of congeneric (or even of closely allied gen- 
era), largely synchronic, monophagous or oligophagous macro-Lepi- 
doptera on a single foodplant species in the North Temperate Zone. 
However, at least eight species of Zale (Noctuidae : Catocalinae) are 
sharing two species of pines in much of southern New Jersey. There 
may well be a comparable assemblage of Catocala on Apocarya in 
the southeastern United States. Comparable assemblages of polypha- 
gous congeners may well occur on some trees. Lithophane would be 
a likely candidate genus, with Quercus spp. and Betula spp. being 
likely hosts. | 


ACKNOWLEDGMENTS 


Lawrence F. Gail accompanied me on several nights of oviposition observations and 
will use some of the data in his own analyses. Thanks are due to him for suggestions 
regarding statistical analyses. David MacDonald took me to the Cheshire and Walling- 
ford localities and assisted with observations there. 


LITERATURE CITED 


FERNALD, M. L. 1950. Gray’s Manual of Botany, 8th ed. American Book Company, 
New York, IXIV + 1632 pp. 

NIELSEN, M. C. 1978. Field Summary for 1977, Michigan Report. News Lepid. Soc., 
No. 3 May/June: 6. 

RICKLEFS, R. E. 1973. Ecology. Chiron Press, Newton, Mass., Portland, Oregon, X 
+ 861 pp. 

SARGENT, T. D. 1976. Legion of Night: The Underwing Moths. Univ. Massachusetts 
Press, Amherst, Mass. XIII + 222 pp. 

1977. Studies on the Catocala (Noctuidae) of southern New England. V. The 
records of Sidney A. Hessel from Washington, Connecticut, 1961-1973. J. Lepid. 
soc. ol: 1=16. 

SCHWEITZER, D. F. 1982. Field observations of divergent resting behavior among 
hickory feeding Catocala larvae (Noctuidae). J. Lepid. Soc. 36(4):303. 


Journal of the Lepidopterists’ Society 
36(4), 1982, 264-268 


THE BUTTERFLIES OF KENT ISLAND, 
GRAND MANAN, NEW BRUNSWICK 


G. DAvID MADDOX! 
Duke University, Department of Zoology, Durham, North Carolina 27705 


AND 


PETER F. CANNELL 


American Museum of Natural History, Department of Omithology, 
79th St. & Central Park West, New York, New York 10024 


ABSTRACT. A list of 23 species of butterflies found to be on Kent Island, New 
Brunswick has been compiled. 


Kent Island, New Brunswick (44°35’N, 66°45’W) is the southern- 
most island in the Grand Manan archipelago in the Bay of Fundy. 
Kent Island is approximately 7.2 kilometers from Grand Manan and 
9.3 kilometers from Maine, which is the nearest point of mainland 
(Fig. 1). 

Thirty-four of Kent Island’s 75.15 hectares are characteristically Ca- 
nadian Zone forest, dominated by white spruce (Picea glauca) and 
balsam fir (Abies balsamea). The rest of the island is open grassy 
fields. McCain et al. (1973) and McCain (1975) have described the 
vegetation in some detail. Kent Island’s weather is dominated by cool 
wet maritime air with frequent and dense summer fog. The Bowdoin 
College Scientific Station maintains a weather recording station on 
the island and data from this station for several meteorological vari- 
ables are presented in Table 1. 

Twenty-three species of butterflies have been found on Kent Island. 
Over 95 species have been recorded in nearby Maine (Brower & Payne, 
1956) and, though it has not been systematically studied, Grand Man- 
an hosts many lepidopteran species not found on Kent Island (Cannell 
& Maddox, personal observations). The number of species occurring 
on Kent Island is probably limited by its remote location, small size, 
and climatic conditions. Gobiel’s (1965) preliminary study of Kent 
Island butterflies included one species, Limenitis archippus Cramer, 
not found during our study, but each of the twenty-three species listed 
here is also found in Maine (Brower & Payne, 1956) and Nova Scotia 
(Ferguson, 1954). 

Many of the species in the following list are known migrants (P. 
interrogationis, V. atalanta, V. virginiensis, N. antiopa, N. j-album, 


’ Present address: Section of Ecology and Systematics, Cornell University, Ithaca, New York 14853. 


VOLUME 36, NUMBER 4 265 


and D. plexippus) and would be expected to reach such an island 
even if they could not breed there. Since there is no Salix or Populus 
on Kent Island L. archippus must be a non-breeding migrant. This is 
surprising since it is usually considered a fairly sedentary species. 

The following list is based on observations conducted between June 
and early September 1979 and between June and mid-October 1980. 
Daily abundances for 1980 were derived from daily estimates of pop- 
ulation sizes (counts of flying adults) and are presented in Fig. 2. 
Larvae were not systematically hunted; so, for most species, definite 
statements about breeding status cannot be made. Nomenclature fol- 
lows Howe (1975). Voucher specimens are held at the Bowdoin Col- 
lege Scientific Station, Kent Island. 


SYSTEMATIC LIST 


Danaus plexippus L. Seen regularly in summer and abundant in fall. 
No Asclepias species are found. 

Cercyonis pegala Fabricius. Infrequent and uncommon in both years. 

Limenitis arthemis Drury. One seen in each year. The principal larval 
food plants (Howe, 1975) are not found. However, Alnus and Sor- 
bus, secondary hosts, are common. 

Limenitis archippus Cramer. Seen rarely in 1960 (Thomas Skaling, 
communication to C. Huntington) and once in 1964 (Gobiel, 1965). 
There is no Salix on Kent Island and therefore L. archippus must 
be a migrant. 

Chlosyne harrisii Scudder. Seen only once, on 8 July 1980. The larval 
food plant, Aster unmbellatus, is uncommon. 

Polygonia interrogationis Fabricius. Common for a brief period in 
July of both years. Adults were always seen near an Alnus bog but 
never near the reported larval food plant, Urtica dioica, which is 
common. 

Vanessa atalanta L. Very common in both years. Adults often seen 
visiting flowers. The larval food plant, Urtica dioica, is common. 
Vanessa virginiensis Drury. Frequently seen in 1979, but less com- 
mon in 1980. Usually found in open field of Achillea millefolium. 
Potential larval food plants are abundant, especially Artemisia, 

Gnaphalium, and Anaphalis. 

Nymphalis antiopa L. Frequently seen in August of both years. Larval 
food plants are not present. 

Nymphalis j-album Boisduval. Seen only once, in late August 1980. 
Larval food plants are not present. 

Nymphalis milberti Godart. Uncommon during July of both years. U. 
dioica, the larval food plant, is common. 


266 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Speyeria cybele Fabricius. Very common from early July until mid- 
August. Larval food plants, Viola cucullata and V. pallens, are com- 
mon. 

Speyeria aphrodite Fabricius. Seen once but not collected on 28 June 
1980. 

Speyeria atlantis Edwards. Rare in both years. 

Lycaena phleaus americana Harris. The first butterfly seen each sum- 
mer. This species was common in June and July of both years. A 
second brood seemed to appear in late August (Fig. 2). The principal 
larval food plant, Rumex, is abundant. 

Lycaena epixanthe Boisduval & Le Conte. In each year ten to twenty 
individuals appeared over a two day period in July. These were 
restricted to a small acid bog. The only reported larval food plant, 
Vaccinium macrocarpum, is not found, but V. oxycoccus is locally 
abundant. 

Glaucopsyche lygdamus Doubleday. This species was seen only once, 
in mid-July 1980. Some of the food plants, Lathyrus and Vicia, are 
found. 

Pieris rapae L. Very common in late summer of both years. Crucifers 
are not numerous, but Cakile edentula and Capsella bursa-pastoris 
are present. August females were seen ovipositing on the latter 
species. 

Colias eurytheme Boisduval. Not seen in 1979 and uncommon in 1980. 
Several Trifolium species are common. C. eurytheme is an annual 
immigrant to this region but is probably unable to overwinter. It is 
surprising that C. philodice, a much commoner insect in New 
Brunswick and Nova Scotia, did not occur on Kent Island. 

Colias interior Scudder. Fairly common in August 1980. Several Vac- 
cinium species are uncommon. 

Papilio glaucus L. Seen once in 1979 and once in 1980. One of the 
larval food plants, Sorbus, is common. 

Papilio polyxenes Fabricius. Seen twice in late August 1980. 

Polites coras Cramer. The only skipper seen in the two years, on 21 
July 1980. The food plant in nature is unknown. 


ACKNOWLEDGMENTS 


We wish to thank Amanda Cannell for advice on aesthetic moralism, Jeff Cherry for 
lively discussions about food and politics, Lisa Reid for optimism, and all of the above 
for help with the collections. We also thank an anonymous reviewer for comments on 
the manuscript and Dr. Charles Huntington and Bowdoin College for our use of the 
Kent Island Scientific Station. Special thanks to Ronny Brite. This is contribution no. 
50 of the Bowdoin College Scientific Station. 


VOLUME 36, NUMBER 4 267 


LITERATURE CITED 


BROWER, A. E. & R. M. PAYNE. 1956. Check list of Maine butterflies. Maine Field 
Naturalist 12:42-44. 

FERGUSON, D. C. 1954. The Lepidoptera of Nova Scotia, Pt. 1, Macrolepidoptera. 
Proc. Nova Scotian Inst. Sci. 23(3):161-375. 

GoBEIL, R. E. 1965. Butterflies on Kent Island. J. Lepid. Soc. 19:181-183. 

Howe, W. H. 1975. The butterflies of North America. Doubleday & Co., New York. 

McCain, J. W. 1975. A vegetational survey of the vascular plants of the Kent Island 
Group, Grand Manan, New Brunswick. Rhodora 77: 196-209. 

, R. B. PIKE & A. R. HODGDON. 1973. The vascular flora of Kent Island, Grand 

Manan, New Brunswick. Rhodora 75:211-—220. 


TABLE 1. Meteorological variables for Kent Island in 1979 and 1980. 


1979 1980 

June July Aug June July Aug 
Mean daily maximum (°C) — 16.9 ies 14.3 17.4 18.2 
Mean daily minimum (°C) — 10.2 10.8 2 9.9 11.4 
Mean daytime (°C) = 14.6 14.7 12.4 14.5 16.6 
Precipitation (cm) — 12.0 12.9 9.7 18.9 U8 
Days with dense fog — 1g W 5 11 ul 
Maximum temperature (°C) — he) 22 20 22 24 


Minimum temperature (°C) — 8 10 3 8 10 


268 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


NEW BRUNSWICK 


45° 


44° 


68° rer 66° | 65° 


Fic. 1. Map of the Bay of Fundy indicating the position of Grand Manan and Kent 
Island. | 


JUNE JULY AUG SEPT OCT 
0 20 oo 18) Zu =a 10 20 30 10 20 0 10 20 30 
D. plexippus ue ‘1 wit u ner sjedii 
P. interrogationis ‘ta8 bh ‘ 
V.atalanta et | tl LU Oe | 
V. virginiensis er ible | 
N. antiopa : ©) sth Be hb Pdi oll ‘ 
N.milberti eon s 
S. cybele OR ee Ee ae 
L. phleas ot tL en | en Per 
P rapae 10 6 ) eae ee | | [aq nal) Bi 
G. eurytheme ' Js ee. 
C. interior : bs pag! ny Hid a 


Fic. 2. Abundance and phenology for 1980 of the eleven species seen at least four 
times in 1980. 


Journal of the Lepidopterists’ Society 
36(4), 1982, 269-278 


FLIGHT PATTERNS AND FEEDING BEHAVIOR OF ADULT 
MILIONIA ISODOXA PROUT AT BULOLO, 
PAPUA NEW GUINEA (GEOMETRIDAE) 


F. R. WYLIE 


Department of Forestry, Biology Section, 80 Meiers Road, 
Indooroopilly, Queensland, 4068 


ABSTRACT. At Bulolo, daily flight activity of adult Milionia isodoxa commenced 
between 0600-0630 h when the moths began moving from hoop pine (Araucaria cun- 
ninghamii Ait. ex D. Don) plantations to adjacent feeding areas (the females to blossoms 
and the males usually to sites along streams and roads). The numbers of males at the 
feeding sites reached a peak between 0800-0915 h and by 1030 h most had returned 
to the plantations; females usually returned towards 1200 h. Field observations sug- 
gested that atmospheric conditions influenced flight behavior of the moths but no direct 
relationship was found. 

Adults feed on floral nectar and the males presumably supplement their diet with 
solutes contained in wet sand, mud, animal dung and carrion. Males preferred to feed 
at the decomposing bodies of the toad, Bufo marinus Linnaeus, and at fresh cattle 
dung. A few males fed at dried toad bodies and at dry dung by moistening the surface 
with a droplet of anal liquid which they then imbibed. Observations suggested that 
odor was important in aggregating males at feeding sites, although some visual stimuli 
may be involved. 


Studies in Papua New Guinea on the taxonomy, distribution and 
biology of Milionia isodoxa Prout, an endemic defoliator of hoop pine 
(Araucaria cunninghamii), were reported by Wylie (1974a, b, 1982). 
Only brief references were made there to studies of adult populations 
of this insect. The majority of adult Geometridae are nocturnal, but 
the adults of M. isodoxa are day flying (Wylie, 1974b) and are rarely 
attracted to light at night. They are seldom found far from where their 
larval host plant, A. cunninghamii, occurs and no long distance mi- 
gratory patterns have been observed. Females feed almost exclusively 
on floral nectar, a habit common in Lepidoptera (Norris, 1936). The 
males, on the other hand, appear to feed primarily on solutes con- 
tained in wet sand, mud, animal dung and carrion, and are frequently 
seen in large groups at such sites. At Bulolo, the feeding sites for each 
sex were usually widely separated (several hundred meters or more) 
and the normal sex ratio of 1:1 was approached only in collections 
made within the plantation canopy (Wylie, 1974b). Studies of the dai- 
ly movement of adults to and from their feeding areas and the feeding 
behavior of the males, carried out at Bulolo during the period April 
1968 to April 1970, are reported below. 


MATERIALS AND METHODS 


Five sites regularly frequented by adult M. isodoxa within and ad- 
jacent to hoop pine plantations were chosen as observation stations. 


270 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Fic. 1. Feeding site (station 2) at Buiolo. Plates of “attractants” used in the food 
preference study are shown. Margin of hoop pine plantation is visible on the left and 
natural forest on the right. 


Stations 1, 2 and 4, each approximately 50 m? in area, were situated 
at nearly 1 km intervals along an unsealed road, 1 and 2 being at 
shallow creek crossings (Fig. 1) and 4 at a cattle grid. These three 
stations were in the open and received direct sunlight during most of 
the day. Station 3 (area 300 m”) extended nearly 50 m along both banks 
of a shallow stream, the margins of which were, in parts, closely 
bounded by secondary vegetation and received patchy sunlight. Sta- 
tion 5 (area 50 m?) was located beneath the plantation canopy (tree 
age eight years at commencement of study) almost equidistant (50 m) 
from stations 2 and 3 and received only diffuse sunlight. Adults gath- 
ered on mud, decaying organic matter, wet sand and rocks at stations 
1, 2 and 3, on cattle dung and mud at station 4 and on hoop pine 
foliage at station 5. | 

In April 1968, the number of adults found at stations 14 were 
counted at 15 minute intervals during daylight hours (0530-1800 h 
approximately) for seven consecutive days. Similar seven-day counts, 
conducted at three monthly intervals from July 1968 to April 1970, 


VOLUME 36, NUMBER 4 271 


were restricted to the period 0700-1100 h, when nearly all adult ac- 
tivity at the stations occurred. Additional counts were made simulta- 
neously at stations 3 and 5 for three days in March 1969 to determine 
flight patterns between the open areas and the adjacent forest. 

In March 1970, a simple experiment was conducted at stations 1, 2 
and 4 to determine the relative attractiveness to M. isodoxa adults of 
five substances: fresh cattle dung, decomposing toad carrion, urine, 
honey-water (1:2 mixture) and water. Five white porcelain dining 
plates (19 cm diameter), each containing one of these substances, and 
two empty plates as controls were positioned at each station with a 
distance of 2 m or more between adjoining plates. The surface area 
of “attractant” on each plate was approximately 200 cm’; the urine, 
honey-water and water were soaked onto pieces of cotton wool to 
provide a footing for the adults. These “attractants’’ were replenished 
daily and their relative positions at the stations rotated to ensure uni- 
formity. Where possible, extraneous attractive material such as carrion 
and dung was removed from the vicinity of the stations at this time. 
Counts were made of the numbers of adults present at each plate at 
15 minute intervals from 0700 h to 1100 h daily for seven consecutive 
days. 

During the studies, bulb thermometers (0—100°C), Lambrect ther- 
mohygrographs® type 252 and solar radiation recorders® model R401 
were used to measure temperature, relative humidity and solar radia- 
tion at the stations. Records of the number of sunshine hours each 
day were obtained from the meteorological station at the Bulolo For- 
estry College, approximately 3 km from the study area. Other details 
recorded included subjective assessments of cloud cover and rainfall 
intensity and observations on the feeding behavior of the males. 


RESULTS 
Flight Patterns 


At the open sites (stations 1-4) the adults were predominantly males, 
and they displayed a similar daily flight behavior with little variation. 
They first appeared at the stations at approximately 0700 h, numbers 
reaching a peak between 0800-0915 h and declining rapidly there- 
after so that by 1030 h few remained at the stations (Fig. 2). Within 
the plantations, adults resting overnight on hoop pine foliage became 
active between 0600-0630 h and began moving to the feeding areas. 
Few adults were seen within the plantations between 0800-0900 h 
each day. Counts at stations 3 and 5 showed that, as the number of 
adults along the stream decreased after 0900 h, numbers within the 
plantations increased, reaching a peak at approximately the same time 


ie JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


fa 
Lu 
Z 
= 
O 
O 
” 
zi 
= 
a 
< 
LL 
oO 
jog 
Lu 
jaa) 
= 
5 
Z 
Zz 
<x 
au) 
= 


Fic. 2. Typical daily flight pattern of male M. isodoxa at feeding sites in the open 
at Bulolo. Means for station 1 are shown for 58 days of observation. 


that the last adults left the stream (Fig. 3). Females observed at blos- 
soms adjacent to the study area continued to feed for some time after 
the last males had been recorded at the stations and returned to the 
plantations towards 1200 h. Both sexes tended to congregate for a time 
on foliage of hoop pine trees along the margins before dispersing 
deeper into the plantation. 

Field observations suggested that adult activity at the stations was 
greater on warm, sunny days, and population peaks occurred earlier 
in the morning than on cool, overcast days. For example, during counts 
at station 2 on 29 March 1969 in cool, overcast conditions (22.2-24.8°C), 
numbers of moths reached a peak of 66 at 0915 h. During counts at 
the same station four days later in warmer, sunny weather (22.7— 
27.8°C), numbers reached a peak of 160 at 0745 h. However, there 
were some cool, overcast days when large numbers of moths and early 
population peaks were recorded and some hot, sunny days when low 
numbers and late peaks occurred. 

At all stations most population peaks (163 or 72% of a total of 225 


VOLUME 36, NUMBER 4 KT) 


ms 
Bee CD number of adults L 
60+ anenie . 39 
Qsseccecees e HE NY pie}rie Cue Res ee ee le ge oe oer 
Decwesesce 
50-4 > 


NUMBER OF ADULTS COUNTED 
LS} _ ~ 
jo) ° ° ° 
It it [Ss 
5 aS | 
° 
@) NI dAHYOALV YHoHd WAL 


+ 20 


ra ne me 


——_= number of adults 


60 FOREST ie 


resconosstecr) temperature 


0800 09 00 1000 1100 
TIME OF DAY 


Fic. 3. Number of adults counted at station 3 (open) and station 5 (forest) between 
0800-1115 h on 5 March 1969. 


observations of peak time) occurred between 0800 and 0915 h each 
day. The earliest peak time recorded was 0730 h and the latest 1015 
h. Of a total of 41 peaks recorded after 0915 h, 21 occurred at station 
3 while, of the remainder, the numbers recorded at stations 1, 2 and 
4 were six, five, and nine respectively. This again suggested that at- 
mospheric conditions influenced flight activity of the moths, since 
station 3 received more shade during the morning than did the other 
three stations. However, peak time of activity at station 3 was not 
consistently later than, and in a few instances preceded peak times 
at the other stations. 

When the data was subjected to stepwise multiple regression anal- 
ysis, no direct relationship was found between adult flight behavior 


274 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


and climatic conditions. The total number of adults (for any particular 
day) and peak time of activity (time during any particular day at which 
numbers reached their peak) varied at random. No definite seasonal 
effect was observed in the numbers of adults counted during the sev- 
en-day study periods in January, April, July and October each year. 


Food Preferences 


The total numbers of adults counted at all stations during the study 
period on dead toad, dung, urine, honey-water, water and controls 
were 1237, 633, 203, 52, 37 and 14 respectively. Analysis of variance 
showed a significant difference at the 1% level between the numbers 
of adults counted at the different “‘attractants.” There was no signifi- 
cant difference at the 5% level between station replications. Numbers 
counted on toad and dung differed significantly from each other and 
from the remaining “attractants” (LSD = 135.7). Although some adults 
fed on wet sand and mud away from the plates, their numbers were 
much fewer than those recorded at toad and dung. 


Feeding Behavior of Males 


Males greatly outnumbered females in collections made on wet 
sand, mud, animal dung and carrion at Bulolo (Wylie, 1974b). Many 
of these moths appeared to be recently emerged; their wing and body 
scales were intact and there was no sign of the wear usually apparent 
on older insects. They probed and fed (with the proboscis extended) 
at moist sand and mud along stream and puddle margins, and occa- 
sionally at moss on rocks and at stream debris, not at the free water 
itself. On very wet ground the moths perched on the edge of dry 
pebbles and probed at the mud below. Several fed at patches of ground 
moistened by human urine and some were attracted to human sweat 
on both clothes and body. Males also fed on floral nectar and, in the 
laboratory, on a honey-water solution. When feeding, the males were 
remarkably docile and were easily captured by hand. 

Large numbers of males fed commonly at cattle dung and toad (Bufo 
marinus) carrion. Fresh dung was preferred to dry, but toads dead for 
at least a day and beginning to decompose appeared more attractive 
than recently killed or dry material. A number of moths observed at 
dry dung and dry toad carcasses exuded a drop of liquid from the anus 
and imbibed from the moistened surface. This behavior among Lep- 
idoptera has previously been regarded as a habit confined to the Hes- 
periidae (Norris, 1936). 

It is interesting to note that while male M. isodoxa feed at the dead 
bodies of B. marinus, they are themselves prey to the toad, as evi- 
denced in the laboratory when a supposedly dead toad revived among 


VOLUME 36, NUMBER 4 275 


captive moths, and in the field, when males attracted to a dead toad 
in a culvert were eaten by other toads sheltering in the culvert. 


DISCUSSION 


There are few references in the literature to adult behavior of Mil- 
ionia species. De Mesa (1934, 1938) records daily flight patterns sim- 
ilar to those of M. isodoxa for adults of the pine-needle measuring 
worm Milionia coronifera Swinhoe at Baguio in the Philippines. He 
notes that adult M. coronifera migrate to the flowering plants between 
0600-0900 h, returning to the pine forests (Pinus kesiya) between 
0900-1000 h each day. A second migration to blossoms occurs at 1700 
h, and the moths rest at night in the crowns of the pine trees. At Bulolo 
there was no conspicuous late afternoon migration of M. isodoxa, al- 
though occasional moths were seen in flight along the roads at this 
time. In Sumatra, Mangundikoro and Depari (1958) mention only that 
moths of Milionia basalis Walker, a defoliator of Pinus merkusii, were 
frequently observed in flight during the day along the fringes of the 
plantations. 

Eight species of Milionia, in addition to M. isodoxa, have been 
recorded from the Bulolo/Wau area. Two of these species, M. dohertyi 
Rothschild and M. mediofasciata Rothschild, are predominantly night- 
flying and regularly appear in collections at light. No daylight flight 
has been observed for moths of M. dohertyi (J. J. H. Szent-Ivany, 
pers. comm.) but a single M. mediofasciata male was recorded at 
station 1 at 0745 h during counts in January 1970. Both M. callima 
Rothschild & Jordan and M. grandis adults were observed in morning 
flight near the plantations, and M. grandis was also collected at light. 
The remaining four species, M. ? aglaia Rothschild & Jordan, M. 
aroensis Rothschild, M. diva Rothschild & Jordan and M. paradesia 
Jordan, were captured in flight during the day along mountain streams 
(1700 m) near Wau (P. Shanahan, pers. comm.). 

Although the daily flight patterns of adult M. isodoxa at Bulolo, 
involving a movement from the plantations to obtain food and a sub- 
sequent return to mate and egg-lay, are established, the factors influ- 
encing the timing of these flights to and from the feeding areas remain 
unknown. Initial observations suggested that either temperature or 
solar radiation may be important in determining when numbers peak 
and when the moths begin to leave the stations. These factors could 
act directly by heating the body of the insect, or indirectly by drying 
out the food and rendering it less attractive. However, this does not 
explain the sometimes early disappearance of moths from shaded areas 
along creeks or roads or the rapid decline in numbers at the stations 
on some cool, overcast days. Similarly, relative humidity or saturation 


216 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


deficiency must be of minor importance when considering a stream 
environment even on the hottest days. 

Rainfall may deter or delay the flight of moths to the stations on 
certain days. It may also affect the numbers of moths recorded at the 
stations on subsequent fine days by (i) increasing the number of al- 
ternative feeding sites and thus decreasing totals at the stations or (ii) 
increasing the attractiveness of the stations as feeding sites and thus 
increasing total numbers recorded. However, it would be unlikely to 
affect flight times on these fine days. 

Many records in the literature indicate that atmospheric conditions 
are important in influencing flight patterns of puddle-frequenting 
species (see Norris, 1936). However, because of anomalies such as 
those described above for M. isodoxa, a simple relationship appears 
unlikely. 

Norris (1936), in an extensive review of the feeding habits of Lep- 
idoptera, lists many species which feed at puddle margins, dung and 
carrion, including some Geometridae. Nearly all records show a great 
preponderance of males at sites similar to that found for M. isodoxa 
at Bulolo. She notes that, as with M. isodoxa, many species drank at 
puddle and riverside sites contaminated by animal excreta even when 
cleaner water was abundantly available and observes that practically 
all water-drinking may be primarily due to the attraction of such con- 
taminants. Downes (1973) concurs, regarding puddles as areas of con- 
centration of organic debris and solutes and further suggests that pud- 
dles, dung and carrion represent successively higher levels of the 
same stimuli. Observations of M. isodoxa at Bulolo support this view, 
toad carrion and dung in that order being preferred to wet sand and 
mud. 

With respect to carrion and dung and perhaps other foods, there 
appear to be different degrees or stages of attractiveness. For example, 
Payne and King (1969) list 21 species of Lepidoptera attracted to pig 
carrion at Clemson, South Carolina, 17 of which (including three geo- 
metrids) preferred the carrion in a state of advanced decay. At Bulolo, 
M. isodoxa preferred toads which were actively decomposing to fresh- 
ly killed or dried material, and more were attracted to fresh dung than 
to dry. 

The nutritional requirement for insects that is satisfied by probing 
at puddles, dung and carrion is little known (Downes, 1973; Norris, 
1936). In Diptera, protein is required by some species for reproduc- 
tive development in females. For example, the females of the sheep 
blowfly, Lucilia cuprina (Wiedemann), require protein to mature each 
cycle of oocytes (Williams et al., 1977), and in many species of mos- 
quitoes the number of oocytes that reach maturity is related to the 


VOLUME 36, NUMBER 4 WIT 


amount and type of blood that the females ingest (Shelton, 1972). In 
Lepidoptera, males of most species are seen puddling much more 
often than females. Puddling behavior probably permits them to take 
in nutrients above those provided by larval nutrition or available from 
nectar, and both organic and inorganic solutes may be sought (Downes, 
1973; Arms et al., 1974). 

The tendency for many puddle-frequenting Lepidoptera to gather 
in closely packed groups at the feeding sites is often mentioned in 
the literature (Norris, 1936). Collenette and Talbot (1928) show how 
feeding aggregations of Catopsilia (Pieridae) are built up by visual 
recognition of the already established individuals. Downes (1973) 
however, suggests that odor may be more important than visual stim- 
uli for some species, although visual responses may also occur. In the 
case of M. isodoxa at Bulolo, odor seems to be of major importance 
in the formation of feeding groups. The most striking aggregations of 
moths are seen on carrion and dung, while they tend to feed more 
individually at lightly contaminated and less odorous areas such as 
stream margins. However, several males, on arriving at the stations, 
flew directly to join groups feeding on toad and dung, a behavior 
which suggests some visual response. 

The described flight behavior of M. isodoxa adults may influence 
the distribution of larval populations of the insect in hoop pine areas. 
At Bulolo, Wylie (1982) showed that M. isodoxa larvae were more 
abundant on host trees in areas adjacent to a stream or soak than on 
trees in drier areas of the plantations. This was attributed, in part, to 
a presumed higher frequency of egg-laying where adults congregate 
on trees close to the feeding sites. 


ACKNOWLEDGMENTS 


I thank Dr. B. Gray, Beerwah, Queensland and Drs. G. Hooper and T. Woodward, 
University of Queensland, Brisbane, for their helpful advice during the study and in 
the preparation of this paper. I thank Dr. D. Sands, C.S.I.R.O., Brisbane for his review 
of the final manuscript. Appreciation is expressed to the staff of the Entomology Sec- 
tion, Bulolo, who assisted in the field studies. I am grateful to Mr. M. Nester, Depart- 
ment of Forestry, Brisbane, for his assistance in the statistical analysis and Mrs. J. 
Hockey and Mrs. D. Toomey for typing the manuscript. 


LITERATURE CITED 


ARMS, K., P. FEENY & R. C. LEDERHOUSE. 1974. Sodium: Stimulus for puddling 
behaviour by tiger swallowtail butterflies, Papilio glaucus. Science 185:372-374. 

COLLENETTE, C. L. & G. TALBOT. 1928. Observations on the bionomics of the Lep- 
idoptera of the Matto Grosso, Brazil. Trans. Entomol. Soc. London 76:391-414. 

DE MgsA, A. 1938. Pine-needle measuring worm Milionia coronifera Swinhoe. Phil- 
ipp. J. Forest. 1:3-13. 

DOwneEs, J. A. 1973. Lepidoptera feeding at puddle-margins, dung, and carrion. J. 
Lepid. Soc. 27:89-99. 


278 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


MANGUNDIKORO, A. & K. S. DEPARI. 1958. Gungguan hama dihutan Pinus merkusii 
di Sumatra utara. Rimba, Indonesia 7:417—-452. 

Norris, M. J. 1936. The feeding habits of the adult Lepidoptera Heteroneura. Trans. 
Roy. Entomol. Soc. London 85:61-90. 

PAYNE, J. A. & E. W. KING. 1969. Lepidoptera associated with pig carrion. J. Lepid. 
Soc. 23:191-195. 

SHELTON, R. M. 1972. The effects of blood source and quantity on production of 
eggs by Culex salinarius Coquillett (Diptera: Culicidae). Mosq. News 32:31-37. 

WILLIAMS, K. L., L. BARTON BROWNE & A. C. M. VAN GERWEN. 1977. Ovarian 
development in autogenous Lucilia cuprina in relation to protein storage in the 
larval fat body. J. Insect Physiol. 23:659-664. 

WyLiE, F. R. 1974a. Description of stages of Milionia isodoxa Prout (Lepidoptera: 
Geometridae) a defoliator of hoop pine in Papua New Guinea. Bull. Entomol. Res. 
63:641-648. 

1974b. The distribution and life history of Milionia isodoxa Prout (Lepidop- 

tera: Geometridae) a pest of planted hoop pine in Papua New Guinea. Bull. Ento- 

mol. Res. 63:649-659. 

1982. Studies of larval populations of Milionia isodoxa Prout (Lepidoptera: 

Geometridae) in the Bulolo-Wau hoop pine plantations in Papua New Guinea. 

Qld. Dep. For. Tech. Paper No. 31 (in press). 


Journal of the Lepidopterists’ Society 
36(4), 1982, 279-289 


REDISCOVERY OF THE TYPE OF PAPILIO PHINEUS 
CRAMER AND ITS BEARING ON THE GENERA 
PHEMIADES HUBNER AND PROPERTIUS 
EVANS (HESPERIIDAE) 


RIENK DE JONG 


Rijksmuseum van Natuurlijke Historie, Leiden, The Netherlands 


ABSTRACT. The type of Papilio phineus Cramer has been rediscovered. It is shown 
that Hubner’s inclusion of the species in his genus Phemiades and Scudder’s desig- 
nation of it as type-species of Phemiades are based on misidentifications. Evans mis- 
identified a yet undescribed species as Papilio phineus Cramer and erected the genus 
Propertius with type-species Hesperia propertius Fabricius, which is congeneric with 
the true Papilio phineus Cramer. The resulting nomenclatorial mess can be solved by 
designation of the yet undescribed Phemiades species as type-species of Phemiades. 
This species is named Phemiades pseudophineus here. The inclusion of Papilio phi- 
neus Cramer in Propertius results in the sinking of Phemiades albistriga Tessmann 
into synonymy with Phemiades phineus (Cramer) on subjective grounds. 


In a recent paper in this journal, Clench and Miller (1980:107) ex- 
pressed the hope that more of Cramer’s species could be determined 
soon. At least for one species their hope seems to have become a 
reality. Previously, the uncertainty as to the identity of the species 
was due to Cramer's poor descriptions and the unreliability of his 
data and figures. The confusion around the species being dealt with 
here has been caused by mistakes of subsequent authors, to put it 
kindly. The name Papilio phineus seems to have cast a spell on peo- 
ple somehow, as almost every subsequent author added some new 
confusion. Now we face a number of nomenclatorial problems that 
would have been quite unnécessary if only these authors had better 
perused the literature and better corrected their proof prints. It may 
be pardonable to some extent if the mainly English speaking authors 
who dealt with the species had problems in understanding relevant 
Dutch, French and German texts, but it is inexcusable that they ap- 
parently never had them properly translated. In the following I will 
indicate the pertinent errors in the literature and suggest some cor- 
rections, hoping that the name Papilio phineus will not bewitch me 
as well. 

I am grateful to Mr. R. I. Vane-Wright of the British Museum (Nat- 
ural History), London, for the opportunity of studying and describing 
relevant material. 


Original Description and Type of Papilio phineus Cramer 


The original (Dutch) description of Papilio phineus by Cramer (1777: 
123; in the index on page 150, spelled “phyneus’’) reads as follows: 


280 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Fics. 1-6. 1, Papilio phineus Cramer, after the original figure; 2, Papilio phineus 
Cramer, type, upper surface; 3-4, Propertius propertius (Fabricius), upper (3) and 


under surface (4); 5—6, Eheuindes aeeUdlanhnatN sp. n., holotype, upper (5) and under. 
surface (6). 


“Fig. E. Phineus. De geele tekening op de onderzyde der vleugelen is 
niet zo schoon van kleur dan van boven, doch voor ’t overige niet 
onderscheiden. Zy word nevens de twee volgende in Surinamen ge- 
vonden.’ Translation: Fig. E. Phineus. The yellow markings on the 
underside of the wings do not have the beautiful color of the upper- 
side, but for the rest are similar. It is found in Surinam, as are the 
next two species. 


Together with the figure of the upperside (pl. 176, fig. E) (repro- 


VOLUME 36, NUMBER 4 281 


duced in Fig. 1), the description is clear enough. In the present con- 
text it is especially important to note the similarity of the markings 
on the upper and underside of the wings as stated by Cramer. 

According to Cramer the specimen from which the illustration of 
Papilio phineus was made, was in the collection of E. F. Alberti, a 
Reverend of the Lutheran Community in Amsterdam. Reverend Al- 
berti died in 1788, and his collection was apparently sold, possibly 
auctioned in parts as was usual in those days. One collector who was 
active in buying parts of other collections was J. Calkoen, who lived 
in Amsterdam and died in 1813 or 1814. His extensive collection of 
insects was in tum auctioned in parts in 1814. Most of it was bought 
by Reinwardt, Director of “’s Lands Kabinet van Natuurlijke Histo- 
rie’ in Amsterdam. At the founding of the Rijksmuseum van Natuur- 
lijke Historie, Leiden, in 1820 by fusion of the “Kabinet” and the 
very large private collection of Temminck, the Calkoen collection 
constituted the basis for the insect collection. This is how some of 
Cramer's type material came to be housed in the Rijksmuseum. 

A badly worn male from the Calkoen collection (Fig. 2) corresponds 
well with Cramer's figure of Papilio phineus except that the head is 
missing. I have little doubt as to its being the actual type. The yellow 
markings of the upper side are repeated on the underside, those of 
the forewings being a little more extensive and a little paler than on 
the upper side, while the yellow band of the upper side of the hind- 
wing has a pale, almost white color on the underside. There is another 
whitish band on the underside of the hindwing from the base to the 
end of vein 8. The presence of this band, although not mentioned by 
Cramer who often omitted details of markings, does not contradict the 
original description of Papilio phineus. 

It must be stressed here that the rediscovery of the supposed type 
of Cramer is not essential for the following lines. Cramer’s description 
alone is sufficient to point out the incorrect statements prevalent in 
the literature. 


Original Description of Phemiades Hubner, 1819, and 
Selection of a Type-Species 


In his well-known “Verzeichnis” Htibner (1819:112) erected the 
genus Phemiades, which he characterized as having “Alle Fluigel oben 
bandartig angelegt, unten nur zerstreut schwarz bezeichnet”’ (all 
wings marked with bands on the upperside, underside only sparsely 
marked with black). The following species were listed by Hubner 
(with the references given by him): 


1208. Phemiades Ephesus. 


282 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


1209. P. Edippus Cram. 366. E. 

1210. P. Epictetus Fabr. Ent. Hesp. 252. Hubn. Urb. vigil. Mys. 
1211. P. Phineus Cram. 63. G. 

1212. P. Augias Linn. Syst. Pap. 257. 


Although the description of the genus is short, its meaning is clear. 
I like to draw special attention to the underside of the hindwing de- 
scribed as sparsely marked with black. If one still doubts the meaning, 
a glance on the underside of the species listed' makes it clear that 
Hubner meant a plain yellow or tawny underside with some black 
shading on the fore- and hindwing or the latter with scattered black 
dots. The only species that does not at all comply with the description 
s “P. Phineus Cramer,’ because the underside of the hindwing is 
quite different. There is one more peculiar aspect to the listing of this 
species by Hubner: the reference to the figure by Cramer does not 
match up. On plate 63G, Cramer (1775) does not depict Papilio phi- 
neus, but shows another South American skipper, Papilio midas Cra- 
mer. This species, currently placed in the genus Bungalotis Watson, 
does not fit the description of the genus Phemiades either. In the 
male (figured by Cramer) the upper side of the wings does not have 
band-like markings (the underside comes closer to Phemiades). 

Whatever Hubner had before him when placing Papilio phineus 
Cramer in Phemiades, it was certainly not Cramer's Papilio phineus. 
This would have been unimportant if Scudder (1875) had not selected 
this same species as the type-species of Phemiades, apparently with- 
out checking Hubner’s reference and without understanding Cra- 
mers description. As a consequence, the type designation is based 
on a misidentification, and in accordance with Article 70 of the In- 
ternational Code of Zoological Nomenclature, the case must be re- 
ferred to the Commission. The Commission can make a choice from 
one of three possibilities: designate as the type-species the nominal 
species actually involved, which was wrongly named in the type- 
designation (does not apply here as the identity of that species is 
doubtful); designate as the type-species the species named by the 
designator, regardless of the misidentification (does not seem to be a 
good choice either because of the discrepancy between Hubner’s de- 
scription of Phemiades and the true Papilio phineus Cramer and this. 
choice would necessitate more changes of names than the third pos- 


1 The species are currently known as follows: P. Ephesus (Hubner described this species only in 1823) = Ampittia 
dioscorides (Fabricius) (Oriental Region); P. Edippus Cramer = Pardaleodes edipus (Stoll) (Afrotropical Region), 
P. Epictetus Fabricius = Anthopthus epictetus (Fabricius) (Neotropical Region), and P. Augias Linnaeus = Telicota 
augias (Linnaeus) (Oriental Region), see Evans (1937, 1949, 1955). For P. Phineus Cramer, see the main text. 


VOLUME 36, NUMBER 4 283 


sibility); or designate as the type-species a species chosen in con- 
formity with the usage of the generic name prevailing at the moment. 


The Genus Phemiades Hiibner in the Literature 


All authors who have mentioned a type-species uncritically ac- 
cepted Scudder’s designation (Watson, 1893; Godman & Salvin, 1900; 
Hayward, 1950; Evans, 1955; Hemming, 1967). The following is a 
sequential history of the genus Phemiades. 

1. Plotz (1883:233) synonymized Phemiades Hubner and Hesperia 
Auct. (which he made a kind of collective genus). 

2. Apart from Papilio phineus Cramer, Watson (1893:104) placed 
Hesperia Utha Hewitson, 1868, in the genus. The latter is currently 
considered a junior synonym of Pyrrhocalles antiqua Herrich-Schaf- 
fer. From Watson's description of the genus it appears that he had 
only P. antiqua at his disposal, as the description does not fit P. phi- 
neus. 

3. Godman and Salvin (1900) placed Hesperia propertius Fabricius, 
1793, with P. phineus in the same genus, Phemiades, and they even 
remarked that both may be the same species. 

4. Schaus (1902) described Phemiades jamaicensis, without indi- 
cation why he placed it in the genus Phemiades. Currently, this name 
is considered to belong to a subspecies of Pyrrhocalles antiqua (Ev- 
ans, 1955) or to a closely related but separate species (Riley, 1975). 
Apparently, Schaus’ allocation was based on Watson’s concept of the 
genus (see above). 

5. Mabille (1904:149) followed Watson (1893) in placing Papilio 
phineus Cramer and Hesperia utha Hewitson in this genus. He added 
Hesperia propertius Fabricius and listed Hesperia memuca Hewitson 
as a junior synonym of the latter (this will be dealt with later). Maybe, 
due to a practice of overlooking things, Mabille ignored the type- 
species designation by Scudder (again mentioned by Watson) and 
remarked in a footnote that phineus probably belonged to another 
genus. 

6. Draudt (1923) did not mention the type-species. He listed the 
following species: propertius Fabricius (=memuca Hewitson), ja- 
maicensis Schaus, phineus Cramer, simulius Druce, and procax sp. 
nov. The last two species were placed in a new genus, Lindra, by 
Evans (1955); for jamaicensis, see 4 above. 

7. Tessmann (1928) described Phemiades albistriga, which he com- 
pared with propertius and considered closely related to this species 
and phineus. 

8. Hayward (1950) mentioned propertius and phineus as belonging 
to Phemiades. 


284 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


9. Evans (1955) listed in addition to phineus as type-species, Au- 
giades pohli Bell, 1932, Trioedusa milvius Mabille, 1904, and Phe- 
miades vergens sp. nov. as the species of the genus Phemiades. For 
Hesperia propertius Fabricius, mentioned repeatedly above, and 
Phemiades albistriga Tessmann, Evans erected the genus Propertius. 

Summarizing, it can be stated that, since Godman and Salvin (1900), 
there has been agreement that Papilio phineus Cramer and Hesperia 
propertius Fabricius are congeneric, except for Evans (1955). It is 
uncertain and seems unlikely that any of these authors ever checked 
and understood the original description by Cramer, but as will be 
shown in the next paragraph, the two species are really closely re- 
lated. 


Propertius Evans, 1955 Versus Phemiades Hubner, 1819 


The genus Propertius was erected by Evans (1955:303) for the 
species Hesperia propertius Fabricius, 1793 (type-species) and Phe- 
miades albistriga Tessmann, 1928. According to Evans’ description 
the genera Propertius and Phemiades differ in the following respects: 

Propertius—“Antennal club long, slender = ¥3 shaft. Unh with al- 
ternating dark red and pale yellow or white bands. Nudum %. F 17 
mm.” 

Phemiades—“Antennal club short, stout = % shaft. Unh ochreous 
with faint yellow spots. Nudum 8/9. F 17 mm.” 

In an additional description on p. 378, however, the antennal club 
of Propertius is said to be %4 of the shaft (it is very difficult to tell 
where the shaft ends and the club begins). Further, in Propertius the 
male is stated to have an inconspicuous narrow, broken stigma, while 
in Phemiades there are separated brands or a broad grey stigma flanked 
by a black patch on either side. I can add an obvious difference in 
the male genitalia: in Propertius the uncus (Figs. 8, 11) ends broadly 
with an upturned apex flanked by two similarly upturned, pointed, 
lateral processes (apparently a formation of the gnathos), while in 
Phemiades (sensu Evans) the uncus tapers and ends simply with a 
small incision (Fig. 14). I entirely agree with Evans that the species 
placed by him in Propertius are generically distinct from those allo- 
cated by him to Phemiades. 

The true Papilio phineus Cramer agrees with the description of 
Propertius. It is not only very similar to the type-species, but it seems 
to be identical with the other species placed in the same genus by 
Evans, viz. albistriga. As I cannot find a difference between albi- 
striga and phineus, the former is sunk as a junior subjective synonym 
of the latter (syn. nov.). 

Externally (Figs. 1-4), the difference between propertius and phi- 


VOLUME 36, NUMBER 4 285 


Fics. 7-12. Male genitalia of Propertius species. 7-9, P. phineus (Cramer), type. 
7, inside of left valve; 8, dorsal view of uncus and tegumen; 9, left lateral view of 
uncus and tegumen. 10-12, P. propertius (Fabricius). 10, left lateral view of uncus 
and tegumen; 11, dorsal view of uncus and tegumen; 12, inside of left valve. 


neus, apart from the color of palpi and head (cf. Evans, 1955:379), is 
mainly the color of the pale bands on the underside of the hindwing, 
being yellow in propertius and white in phineus. The male genitalia 
(Figs. 7-12) differ in the following respects: although the two species 
are of equal size, and tegumen and uncus are also equally large, the 
valve of phineus is 1% times as long as that of propertius, and the 


286 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


dorsal part of the cucullus of the former is more expanded, covering 
a larger part of the costa. For the rest the genitalia are very similar. 

So, the differences between the two species are slight. Both taxa 
are largely allopatric and for that reason could possibly be subspecies 
of a single species, but as both have been recorded from Venezuela 
and E. Peru (Evans, 1955) and the exact distribution areas are poorly 
known, it seems advisable to consider them separate species for the 
moment. 

As said above, designation of the true Papilio phineus Cramer as 
type-species of the genus Phemiades would result in a genus concept 
that is widely different from what was meant by Hubner. Moreover, 
Propertius would become a junior synonym of Phemiades as phineus 
and propertius are certainly congeneric, and Phemiades (sensu Ev- 
ans) would be in need of a new name. Such an action would create 
more instability than the designation of one of the species of Phe- 
miades (sensu Evans) as type-species of Phemiades Hubner. For this 
choice it must also be kept in mind that Evans’ work is far more 
authoritative than any of the other works mentioned. Finally, Evans’ 
concept of Phemiades is not at variance with Hubner’s. The only 
nomenclatorial change needed is a name for the species that Evans 
mistook for Papilio phineus Cramer, this new name being necessary 
anyway. I would propose to the Commission to select this new species 
as type-species of Phemiades. According to Evans it virtually was the 
type-species, and it does not contradict Hubner’s concept of the ge- 
nus. It can be described as follows. 


Phemiades pseudophineus, new species 


External characters (Figs. 5-6). Length of forewing, 6 17.4-17.7 
mm; ¢ 20.9 mm. Male. Upperside dark brown, orange-brown along 
costa of forewing up to apical spots. Forewing with yellow dash in 
basal half of space la, yellow spots in spaces lb, 2 and 3, and orange 
spots in spaces 4, 5 (both very small and inconspicuous), 6-8 and two 
in cell (small and inconspicuous). Hindwing with a central row of 
yellow spots in spaces lc—6, separated by dark brown veins. Fringes 
forewing dark brown, a shade lighter near tornus, hindwing yellow. 
On the forewing inconspicuous dark brown brands: a V-shaped brand 
at the base of space 2, a short brand just below it in space lb, and a 
dot over vein | at the inner corner of the spot in space 1b. Underside 
forewing along costa and apical third, and all of hindwing with a 
peculiar brownish-ochreous color, rest of forewing dark brown; spots 
on underside as on upper side, less conspicuous because of the paler 
ground color. Female. As male, but on upper side, costa of forewing 


VOLUME 36, NUMBER 4 ZO 


Fics. 13-15. Male genitalia of Phemiades pseudophineus sp. n., holotype. 13, in- 
side of left valve; 14, dorsal view of uncus and tegumen; 15, left lateral view of uncus 
and tegumen. 


orange-brown only in basal half, spots in spaces 2, 3, 6, and 7 hyaline, 
spot in space lb semihyaline, other spots very faint or absent. Un- 
derside yellow where the male is brownish-ochreous, spots on hind- 
wing faint. 

Male genitalia (Figs. 13-15). Uncus tapering, bifid at apex. Costa 
of valve slightly swollen and hollowed at apex where it meets the 
upper edge of the cucullus. Latter broad, gently curved, hollowed at 
- inside forming a narrow longitudinal ledge. 

Identification. The new species can be distinguished at a glance 
from Propertius species by the underside of the hindwing, compare 
Figs. 4 and 6. From the other Phemiades species it can be separated 
by the brand on the forewing consisting of three parts (in the other 
species it is single), and by the broad and gently curved cucullus. 

Material examined. Holotype, 6, Chapada (Brazil). Paratypes: 1 
36, Chapada (Brazil); 1 2, Espirito Santo (Brazil). All types in British 
Museum (Nat. Hist.), London. 

According to Evans (1955:380) there should be 2 d¢ and 22 @ in 
the British Museum (under the name of phineus Cramer), but Mr. R. 
I. Vane-Wright of the said museum doubted if there were ever any 
more specimens than the ones listed above. Evans himself apparently 
was not quite sure of the identity of the specimens, as he added a 
note to one specimen reading, “comes nearer phineus Cram. than any 
other known species.” 


288 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Classification, Distribution and Synonymy 


The following classification is essentially that given by Evans (1955) 
with necessary nomenclatorial changes: 


Propertius Evans, 1955:303. Type-species by original designation: Hesperia propertius 
Fabricius, 1793. 

1. Propertius propertius (Fabricius). From Panama and Venezuela through Peru and 
Bolivia to Paraguay, S. Brazil and Argentine. 

Hesperia propertius Fabricius, 1793:325 (“Indiis”’); Hewitson (1869:70); Plotz 
(1882:452). 

Phemiades propertius; Godman and Salvin (1900:529); Mabille (1904:149); Draudt 
(1923:956); Tessmann (1928:127); Hayward (1950:138). 

Propertius propertius; Evans (1955:379). 

Hesperia memuca Hewitson, 1868:37 (no locality stated), 1869:70 (synonymized 
his own species with propertius); Plotz (1882:452). 

Pamphila theodora Ehrmann, 1907:319 (Venezuela), synonymized by Evans (1955). 

2. Propertius phineus (Cramer), comb. nov. Guyanas, Venezuela, Peru. 

Papilio phineus Cramer, 1777:123 (in index on p. 150 spelled “phyneus’’) (Suri- 
nam); Fabricius (1781:132). 

Hesperia phineus; Hewitson (1869:70; Hewitson’s remark on the underside of 
the hindwing of Cramer's species is based on Cramer's fig. 176C that however 
represents Papilio phyllus Cramer as clearly stated by Cramer); Plotz (1883: 
225) 

Phemiades phineus; Watson (1893:104); Godman and Salvin (1900:529); Mabille 
(1904:149); Draudt (1923:956); Tessmann (1928:127); Hayward (1950:138). 
Mention of this combination by Hubner (1819:112), Scudder (1875:247), and 
Evans (1955:380) does not relate to this species. 

Phemiades albistriga Tessmann, 1928:127 (Montealegre, Pachitea, E. Peru), syn. 
nov. 

Phemiades Hubner, 1819:112. Designation of Papilio phineus Cramer as type-species 
by Scudder (1875:247) based on misidentification. New designation suggested to 
Commission: Phemiades pseudophineus sp. n. Trioedusa Mabille, 1904:144. Type- 
species Trioedusa milvius Mabille, sole species included. Synonymized by Evans 
(1955:379). 

1. Phemiades pseudophineus sp. n. Chapada, Espirito Santo (Brazil). 

Phemiades phineus; Evans (1955:380). 

2. Phemiades pohli (Bell). Ecuador, Brazil, Paraguay, Argentine. 

Augiades pohli Bell, 1932:136 (Santa Catharina). 

Ochlodes pohli; Hayward (1950:52). 

Phemiades pohli; Evans (1955:380). 

Ochlodes kohleri Hayward, 1937:94 (Argentine), synonymized by Hayward (1948: 
106). 

Phemiades pohli cidra Evans, 1955:380 (Archidona, N.E. Ecuador). 

3. Phemiades vergens Evans, 1955:381 (Cosnipata, E. Peru). 
4. Phemiades milvius (Mabille). Peru, Br. Guyana, Brazil. 

Trioedusa milvius Mabille, 1904:145 (Brazil). 

Phemiades milvius; Evans (1955:381). 

Phemiades milvius milor Evans, 1955:381 (Yahuarmayo, Peru). 


LITERATURE CITED 


BELL, E. L. 1932. Notes on some American Hesperiidae and descriptions of new 
species (Lepidoptera, Rhopalocera). Bull. Brooklyn Entomol. Soc. 27:131-141. 
CLENCH, H. K.& L. D. MILLER. 1980. Papilio ladon Cramer vs. Argus pseudargiolus 
Boisduval and Deconte (Lycaenidae): A nomenclatorial nightmare. J. Lepid. Soc. 

34:103-119, figs. 1-14. 


VOLUME 36, NUMBER 4 289 


CRAMER, P. [1775]. De Uitiandsche Kapellen voorkomende in de drie Waerelddeelen 
Asia, Africa en America. S. J. Baalde, Amsteldam. Vol. 1, pp. 1-132, pls. 1-84. 

[1777]. Idem. Vol. 2, pp. 1-151, pls. 97-192. 

DRAvuDT, M. 1921-1923. Grypocera. Pp. 833-1012 in A. Seitz. Die Gross-schmetter- 
linge der Erde. Vol. 5. Alfred Kernen, Stuttgart. 

EHRMANN, G. A. 1907. New tropical American Hesperidae. Can. Entomol. 39:317- 
O25: 

EVANS, W. H. 1937. A catalogue of the African Hesperiidae in the British Museum. 
British Museum, London. Pp. i-xii, 1-212, pls. 1-30. 

1949. A catalogue of the Hesperiidae from Europe, Asia and Australia in the 

British Museum (Natural History). British Museum, London. Pp. i-xix, 1-502, pls. 

1-53. 

1955. A catalogue of the American Hesperiidae in the British Museum (Natural 
History). Part 4. British Museum, London. Pp. 1-499, pls. 54-88. 

FABRICIUS, J. C. 1781. Species insectorum. Tom. II. C. E. Bohn, Hamburg and Kiel. 
Pp. 1-517. 

1793. Entomologia systematica. Tom. III. Pars 1. C. G. Proft, Copenhagen. 

GoDMAN, F. D. & O. SALVIN. 1887-1901. Biologia Centrali-Americana. Insecta, Lep- 
idoptera—Rhopalocera. Vol. 2, pp. 1-782. 

HAYWARD, K. J. 1937. Hesperioidea Argentina, 5. Rev. Soc. Ent. Argent. 9:93-100. 

1948. Hesperioidea Argentina, 19. Acta Zool. Lilloana 5:103-112. 

1950. Insecta, Lepidoptera, Hesperiidae, Hesperiinae. Pp. 1-388 in H. R. 
Descole. Genera et species animalium argentinorum. Vol. 2. Guillermo Kraft, Bue- 
nos Aires. 

HEMMING, F. 1967. The generic names of the butterflies and their type-species. Bull. 
Brit. Mus. (N. H.), Entomology. Suppl. 9, pp. 1-509. 

HEwITSON, W. C. 1867-1871. Illustrations of new species of exotic butterflies. Vol. 
4. John van Voorst, London. Pp. 1-114, pls. 1-60. 

1868. Descriptions of one hundred new species of Hesperidae. Part 2. John 
van Voorst, London: Pp. 26-56. 

HUBNER, J. 1816-{1826]. Verzeichniss bekannter Schmettlinge [sic]. Augsburg. Pp. 
1-431. 

1823. Sammlung exotischer Schmettlinge [sic]. Zutrage. Zweites Hundert. 
Augsburg. Pp. 1-32. 

MABILLE, P. 1904. Lepidoptera, Fam. Hesperidae. In P. Wytsman (ed.). Genera in- 
sectorum, Fasc. 17, pp. 1-210, pls. 1-4. V. Verteneuil & L. Desmet, Bruxelles. 
PLOTZ, C. 1882. Die Hesperiinen-Gattung Hesperia Aut. und ihre Arten (Fortset- 

zung). Stett. ent. Zeit. 43:436—456. 

1883. Idem (Schluss). Stett. Ent. Zeit. 44:195-233. 

RILEY, N.D. 1975. A field guide to the butterflies of the West Indies. Collins, London. 
Pp. 1-224, pls. 1-24. 

SCHAUS, 1902. Descriptions of new American butterflies. Proc. U.S. Natn. Mus. 24: 
383-460. 

SCUDDER, S. H. 1875. Historical sketch of the generic names proposed for butterflies. 
Proc. Amer. Acad. Arts Sci., Boston 10:91-—293. 

TESSMANN, G. 1928. Neue Schmetterlinge aus Ostperu. Mitt. Zool. Mus. Berlin 14: 
117-130, pl. 5. 

WATSON, E. Y. 1893. A proposed classification of the Hesperiidae, with a revision of 
the genera. Proc. Zool. Soc. London 1893:3-132, pls. 1-3. 


Journal of the Lepidopterists’ Society 
36(4), 1982, 290-302 


SELECTION OF OVIPOSITION SITES BY THE BALTIMORE 
CHECKERSPOT, EUPHYDRYAS PHAETON (NYMPHALIDAE) 


NANCY E. STAMP! 
Department of Zoology, University of California, Davis, California 95616 


ABSTRACT. Selection of oviposition sites by the Baltimore checkerspot (Euphy- 
dryas phaeton) was examined in a natural population. Females chose leaves larger than 
average. Egg clusters were clumped, with 1% of the available leaves and 3% of the 
available stalks used. The behavior resulting in this non-random pattern is discussed. 


Butterflies choose host plants which promote larval survival and 
avoid those which do not (Wiklund, 1974, 1975; Chew, 1975, 1977; 
Rausher, 1980). Ovipositing females may discriminate among conspe- 
cific host plants either by avoiding eggs on plants or by depositing 
their eggs with those of other females (e.g. Ehrlich & Gilbert, 1973; 
Gilbert, 1975; Benson et al., 1975; Rothschild & Schoonhoven, 1977; 
Rausher, 1979). Most butterflies deposit their eggs singly (Stamp, 1980), 
but the Baltimore checkerspots (Euphydryas phaeton Drury: Nym- 
phalidae) lay clusters of eggs and tend to deposit egg clusters with 
clusters already present. The advantages for a female in discriminat- 
ing among conspecific host plants and depositing eggs with other egg 
clusters have been linked to avoidance of parasitoids and predators 
and to aspects of the host plants such as the size of the host plant, the 
part of the plant used by larvae, and the distribution of the host plant 
(Benson, 1978; Stamp, 1980). 

My objective was to examine selection of oviposition sites by E. 
phaeton by determining searching behavior, characteristics of the ovi- 
position sites, characteristics of the host plants available for oviposi- 
tion, and distribution of egg clusters. 


METHODS 


E. phaeton was studied at the Conservation and Research Center 
of the National Zoological Park at Front Royal, Warren Co., Virginia 
from 1977 through 1979. This butterfly is univoltine and deposits large 
clusters (x = 274 eggs per cluster; Stamp, 1982c) in June. The early 
instars make communal webs on their larval host plants and then 
diapause in webs in August. The caterpillars overwinter on the ground 
in the plant litter (Bowers, 1978). The larval host plant, turtlehead 
(Chelone glabra L.: Scrophulariaceae), is a clonal perennial growing 
in dense patches (up to 2.3 m in diameter) in wet meadows. Although 


VOLUME 36, NUMBER 4 291 


a few isolated plants consist of one stalk, most plants contain numer- 
ous stalks. 

Plants were searched for egg clusters, and those stalks with clusters 
were tagged. Host plant and leaf-searching behavior were recorded 
by following females searching for oviposition sites. By making quick 
sketches of plants, stalks and leaves and numbering them in order of 
visitation, I kept track of females searching plants, stalks and leaves 
they had previously encountered during the observations. 

In 1979, 107 female E. phaeton were marked using Testor’s enamel 
paint on the wings, thorax and abdomen with no detrimental effects. 
One area (12 x 55 m) was surveyed for host-searching and ovipositing 
females from 1000 to 1700 hours on 9 through 23 June. Each survey 
required about 45 min. Upon finishing a survey, another survey was 
begun immediately. The position of egg clusters was marked with a 
spot of permanent black ink on the upper surface of the leaf and the 
stalk tagged with designation of which female laid the cluster, date, 
height of the cluster from the ground, and leaf upon which the cluster 
was deposited. To determine the proportions of ovipositions actually 
observed, all leaves were examined twice a week for additional egg 
clusters, and those stalks with clusters were tagged. To determine if 
any egg clusters were missed, the area was searched every week 
through the second week of July for untagged stalks with red egg 
clusters (deposited prior to 24 June) and webs of E. phaeton larvae. 
Based on 267 egg clusters (of which 99% were located prior to hatch- 
ing), I observed 87% of the ovipositions in this area over the 14-day 
period using the survey method described above. 

To determine the size of stalks available to E. phaeton, 15 plants 
were sampled. A rod was placed through each plant until 20 stalks 
were partitioned. Those stalks were measured for height and for length 
of leaf nearest to the midpoint of the stalk. The total number of stalks 
and the number of stalks in the outer 15 cm of each plant group were 
counted. The width and perimeter of the plant groups were measured. 
In one area turtlehead was mapped using a reel tape stretched across 
a grid (of 3 X 3m units) and moved at 30 cm intervals along the grid. 
The perimeter of the plant groups and mean width of the plant groups 
were calculated using a map meter (which determined distance). 


RESULTS 
Oviposition Behavior 


Females searching for oviposition sites flew from stalk to stalk about 
0.6 m above the ground along edges of host plant groups, in contrast 
to non-searching females which generally flew above the vegetation 


292 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


TABLE l. Searching behavior exhibited by female E. phaeton (n = 21). 


Mean per female 


based on total Standard 

Searching behavior observations error 
Minutes observed 11.4 ae 14) 
Plant groups visited 1.6 +0.2 
Stalks visited 8.1 +0.9 
Leaves touched 7.5 ert 
Leaves examined Del +0.3 
Turns on top surface of leaves 4.9 +0.8 
Times abdomen placed under leaves 9.2 22117) 
Times returned to top surface of leaves 8.1 a2 16 


without landing on turtlehead. Usually a female settled at a host plant 
after landing on one or two plants and stalks and began oviposition 
after examining an average of two leaves (Table 1). Frequently, a 
female returned to a host plant, stalk or leaf which she had already 
visited during this searching period; 28% of the leaves and 32% of 
the stalks visited had been examined a few to 35 min earlier. How- 
ever, Oviposition on these re-examined leaves was similar to that on 
newly-examined leaves (8 ovipositions on 47 leaves and 10 oviposi- 
tions on 120 leaves, respectively; x?-test, P > 0.10). Oviposition on re- 
examined stalks was also similar to that on newly-examined stalks (9 
ovipositions on 49 stalks and 9 ovipositions on 103 stalks, respective- 
ly; x?-test, P > 0.10). Thus, females appeared to be sampling available 
oviposition sites rather than just responding to each potential site, 
positively or negatively. In addition, females examined and ovipos- 
ited on damaged leaves, partially eaten by sixth instar larvae of E. 
phaeton. 

A female examined a leaf in detail by repeatedly walking over its 
upper surface, tapping it with her antennae, occasionally making 180 
or 360° turns on the leaf and examining the under surface of the leaf 
with her ovipositor while hanging onto the edge of the leaf with the 
midlegs (Table 1). Often a female examined the under surface of a 
leaf by hanging first from one side of the leaf and moving along the 
edge. Then she retumed to the top surface, walked to the other side 
of the leaf, and examined the under surface of the leaf from that side 
(Table 1). 

Each of the 21 females followed oviposited within 22 min. The 
majority of these females were initially flying quickly from patch to 
patch of turtlehead and basking intermittently, rather than having 
started to search among the host plants. Since some other marked 
females were observed to search for up to three hours before ovipos- 
iting, 22 min is a conservative (short) estimate of average search time. 


VOLUME 36, NUMBER 4 293 


TABLE 2. Affect of inclement weather on oviposition. Sixteen and 15 females de- 
posited single clusters and 23 and 22 females deposited eggs with others, before and 
after poor weather, respectively. 


Other clusters 


Ovipositions Single clusters present Total 
Day before: 
Observed 16 15 31 
Expected 16.1 14.9 
Day after: 
Observed 26 24 50 
Expected 25.9 24.1 
42 39 81 


Most females oviposited in early afternoon, although some were ob- 
served ovipositing from 1100 to 1700 hours. Recently-emerged fe- 
males (with bright, unworn wings) and older females deposited clus- 
ters in the moming and afternoon at a similar rate (for recently-emerged 
females, 27 deposited eggs in the moming and 44 in the afternoon; 
for older females, 16 laid eggs in the morning and 62 in the afternoon; 
x?-tests, P > 0.10). Poor flight conditions (e.g., cool, windy, rainy) in- 
hibited egg depositions (x?-tests, P < 0.001; Table 2). However, nei- 
ther the numbers of egg clusters deposited nor the number of females 
depositing eggs was significantly different between the day before 
and the day after inclement weather (y?-tests, P > 0.10). 

Deposition of an egg cluster took an hour and a half (n = 24, x = 
88 min + 37 S.D.). The number of egg clusters laid over a 13-day 
period (by females marked on the first to eighth day of that period) 
ranged from 0 to 6 (n = 98 females, x = 1.3 clusters + 14 S.D.). How- 
ever, the count of egg clusters per female was conservative, because 
13% of the ovipositions in this area were not observed, and some 
females may have oviposited in adjacent areas (about 20 m away). 
Thus, based on a mean of 274 eggs per cluster, a female may lay up 
to 1650 eggs during her lifetime of two to three weeks. Of the 153 
observed ovipositions by marked females, 13% involved females mak- 
ing two or more depositions in a single day. 

Females frequently chose stalks and leaves which already had egg 
clusters (Fig. 1). At the peak of the oviposition period for an area with 
plantain (Plantago lanceolata L., an alternate host plant; Stamp, 1979) 
and by the end of the oviposition period in areas with turtlehead, over 
23% of the stalks with egg clusters had two or more clusters. However, 
of those turtlehead stalks with clusters just after the first week of 
oviposition (n = 46), 47% had two or more clusters. Additional clus- 
ters were deposited a few days apart (2.4 days + 0.3 S.E., range 0 to 


2.94 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Z 40 95 

oS 

w 169 23 
os 136 

@ 20 

oO 

i 

c 

o 

Oo 

= 

o 

a. ~ - 

oe 109 

=3 

c 

o ® 

Oo, 149 210 
ee qe 

eo 

: ; me on 


1977 1978 1979 
Turtlehead Plantain 


Fic. 1. For those stalks and leaves with egg clusters, the percentages of stalks and 
leaves with two or more egg clusters are shown. Numbers above the bars indicate total 
stalks and leaves with egg clusters. Plantain data are from Stamp (1979). 


16 days, n = 82). Of stalks with two or more clusters (n = 42), the 
mean number of depositions was 3.0 (+0.3 S.E.), with up to a total of 
10 clusters per stalk. 

Females laid their clusters with those of other females. Of 153 ob- 
served ovipositions of marked females, 46 females laid eggs with clus- 
ters deposited by other marked females. Only one female laid two 
clusters on the same stalk. For these 153 females (which used 125 
stalks), the binomial probability of two clusters deposited by a female 
in one place is less than 0.001. Twice I observed a female examining 
a leaf on which another female was ovipositing. The searching female 
examined the under surface of the leaf from both sides, crawled across 
and jostled the ovipositing female, and within a few minutes ovipos- 
ited next to the other female with their wings touching. 

Once oviposition began, females exhibited some degree of site te- 
nacity. I observed 10 ovipositing females which were interrupted by 
males or my movements and then flew 1.5 m or more from their clus- 
ters. Within 2 min, three resumed oviposition on the leaves with their 


VOLUME 36, NUMBER 4 295 


TABLE 3. Comparison of oviposition sites among years. One standard error is indi- 
cated and sample sizes are in parentheses. The first four variables were tested each 
with one-way ANOVA; ,x?-test was used for egg clusters in the outer portion of plant 
groups. 


Statis- 
tical 
signifi- 
1977 1978 1979 cance 


Height of stalk incm 74.9 + 1.0(165) 60.7 + 1.1(168) 60.9 + 1.3(166) P< 0.01 
Height of egg cluster 

from groundincm 49.8 + 0.9(165) 43.2 + 0.8(168) 50.8 + 0.3 (166) P< 0.01 
Width of plant 

group in cm 5APATENS.O) CHA) 50:9)=E 1225) 29) 66:95 4-9:(122) P< 0.01 
Leaf length in cm AO OD) s lease 0.2) (68) 0:7 = Ol210168) 0:01 
Percentage of egg 

clusters in outer 

15 cm of width of 

plant group 99.3 (165) 91.7 (168) 90.0 (240) P< 0.01 


partially laid clusters. Within 5 min three others resumed oviposition 
on leaves within 15 cm of the first oviposition sites. The rest of the 
females left the plant groups and did not return that day. 


Oviposition Sites 


Considerable among-year variation occurred in the characteristics 
of oviposition sites: height of stalks chosen by females, height of egg 
clusters from the ground, width of plant groups, and length of leaves 
with clusters (Table 3). This may reflect variation among years in the 
growth of the host plant, probably a consequence of the amount of 
spring precipitation (Stamp, unpubl. data). Most of the egg clusters 
occurred in the outer 15 cm of the plant groups. Females chose larger 


TABLE 4. Dispersion of egg clusters on host plant stalks, based on an estimate of 
6554 available stalks. 


Number of 
clusters Number of Percentage of 
per stalk stalks observed egg clusters 


SCOMOHADOUHWNEH © 
bo 
KF OrFFNOK CO CC 
S 


= 


296 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


n= 238 300 


Height of stalk 
in cm 


Leaf length 
in cm 


Stalks with Sampled stalks 
egg clusters 


Fic. 2. Comparison of oviposition sites and stalks available for oviposition in 1979, 
with + one standard error. Numbers above bars are sample sizes. 


stalks and larger leaves than the mean available (two-sample t tests, 
P < 0.01 for both variables; Fig. 2). Stalks with multiple clusters were 
compared to stalks with solitary clusters deposited late in the flight 
period to evaluate the hypothesis that multiple clusters occurred on 
larger leaves and stalks. However, the height of stalks and length of 
leaves with eggs were similar for stalks with solitary and multiple 
clusters (x = 59.6 cm + 2.6 S.E. and x = 62.3 cm + 4.6 S.E. for stalks, 
and x = 11.0 cm + 04 S.E. and x = 11.2 cm + 0.6 S.E. for leaves, 
respectively; n = 22 solitary and 25 multiple clusters, two-sample t 
tests, P > 0.50 for both variables). E. phaeton used 53 to 59% of the 
plant groups available to them. Also, plant groups were classified by 
size: perimeter less than 100 cm, between 100 and 500 cm, and greater 
than 500 cm. These butterflies were not selecting plant groups by size 
(4 of 17, 34 of 58 and 7 of 7 of the plant groups had egg clusters for 
the respective plant group sizes, y?-test, P > 0.05). In 1977 and 1979, 
egg clusters were randomly distributed among the quarters of plant 
groups (x?-tests, P > 0.05). However, in 1978 the southwest quarter 


VOLUME 36, NUMBER 4 297 


TABLE 5. Dispersion of egg clusters on leaves, based on an estimate of 26,216 
available leaves. 


Number of 
clusters Number of Percentage of 
per leaf leaves observed egg clusters 
0 26,006 — 
1 ar 68.3 
% 25 19.3 
3 3 Si) 
4 Y Oo 
5 3 5.8 


of plant groups had significantly more egg clusters (P < 0.01). This 
was probably a consequence of the availability of edges of host plant 
groups, with some edges not discovered by butterflies due to the 
height of adjacent vegetation. 

I estimated the number of turtlehead stalks and leaves available to 
E. phaeton for oviposition in one area, based on stalks in the outer 
15 cm of the plant groups and large leaves on the upper half of stalks. 
Although these stalks and leaves are referred to as available, no as- 
sumption is made here that they are necessarily suitable to ovipositing 
females. A mean of one stalk per 3.8 cm + 0.2 S.E. of perimeter was 
calculated. This mean was multiplied by the total perimeter of the 
mapped plant groups to estimate the number of available oviposition 
sites. The estimate was 6554 stalks available to E. phaeton for ovi- 
position in this area. The estimate of number of leaves available for 
oviposition was calculated based on four large leaves on the upper 
half of the stalk for each stalk in the outer 15 cm of the plant groups 
rather than for all leaves (mean of 18) per stalk. I multiplied four 
leaves per stalk by 6554 stalks to obtain an estimate of the number of 
leaves per stalk which were available for oviposition. This estimate 
was 26,216 leaves. 

Only 2.6% of the estimated available stalks and 0.8% of the esti- 
mated available leaves were used by E. phaeton for oviposition (Ta- 
bles 4 and 5). These values may be lower if more of the stalks and 
leaves were available as oviposition sites. Females did occasionally 
use stalks near the center of the plant groups as well as leaves on the 
lower half of stalks. The index of dispersion was used to determine 
if egg clusters were distributed randomly on stalks and leaves (South- 
wood, 1978). The number of clusters per stalk and per leaf indicated 
clumped distributions significant at the 0.001 level. 

Some stalks may provide more stimulus for oviposition than others 
and, consequently, E. phaeton may respond to these stalks rather than 
to the egg clusters on them. At four-day intervals from 13 through 25 


298 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Number of clusters 


ine) 
‘ ee | itil : 


Stalk 


Fic. 3. Repeated use of particular stalks for egg deposition in a moderate-sized 
plant group. Leaves with egg clusters were removed from these stalks on each date. 
Total number of leaves with clusters are indicated above bars. No clusters were found 
on 25 June, at the end of the flight period. 


June, the stalks in one plant group were checked for clusters and 
leaves with clusters were removed. The plant group was 74 cm in 
diameter with more than 30 stalks in the outer 15 cm of the plant 
group. Over a period of 12 days, six stalks had clusters (Fig. 3) and 
the mean number of clusters on those stalks was 3.7 (+2.7 S.D.). Thus, 
females were repeatedly using particular stalks even though I was 
removing leaves and the potential stimulus of those stalks either due 
to those leaves or the egg clusters was reduced. 


DISCUSSION 


Overall, E. phaeton females carefully assessed the suitability of 
host plant groups and stalks by detailed examination of the leaves 
upon which they oviposited. E. editha are also known to spend con- 
siderable time in search of oviposition sites and in depositing egg 
clusters (Labine, 1968). E. phaeton do not appear to be discriminating 
among plants to avoid parasitoids, predators or competitive larvae. 


VOLUME 36, NUMBER 4 299 


Egg parasitoids and predators are of more immediate concern than 
larval enemies, because the eggs are exposed on the leaves three 
weeks before hatching. However, E. phaeton oviposited frequently 
on leaves and stalks that they examined previously a few seconds to 
many minutes earlier. This suggests that females were responding to 
other qualities of the stalks and leaves than the presence of egg para- 
sitoids and predators. The sequence of egg clusters deposited on a 
leaf did not affect the level of parasitism or number of parasitized 
eggs per cluster (Stamp, 1981b). Furthermore, loss of eggs to predators 
was small, with no clear difference between single and multiple clus- 
ters on leaves (Stamp, 1981b). Thus, depositing eggs with those of 
other females did not lower the risk to eggs, as it might by surrounding 
eggs with others, predator satiation, or causing parasitoids to run out 
of their own eggs. 

E. phaeton chose larger leaves on larger stalks than those generally 
available and only used a small portion of the available host plant. 
Furthermore, females oviposited repeatedly on particular stalks, sug- 
gesting a paucity of attractive stalks. Perhaps these butterflies were 
selecting by chemical cues particular stalks of plant groups which 
promote higher survival of offspring and consequently, yielded a 
clumped distribution of egg clusters. The fact that more stalks re- 
ceived multiple depositions early in the flight period than later sug- 
gests that females later in the flight period may have had to choose 
between particularly attractive stalks (frequently with several egg 
clusters) and less attractive -stalks. The clonal, perennial turtlehead 
may benefit by producing a few attractive stalks if that reduces the 
number of potential flowering stalks which are destroyed by this her- 
bivore. This would be similar to poplar (Populus angustifolia) pro- 
viding a limited amount of optimal resources and thereby, restricting 
successful colonization by a gall-making aphid (Whitham, 1978). 

Frequently E. phaeton deposited their eggs with those of other 
females. Depositing eggs with other clusters has been observed in 
other populations of E. phaeton (Bowers, 1979) and in populations of 
E. gillettii (Williams, 1981) and E. aurinia (Keith Porter, pers. comm. ). 
The proposition that E. phaeton may deposit eggs with those of others 
with the result that larvae benefit from large group size (from two or 
more egg clusters) was examined in detail (Stamp, 1981a, b, 1982a, c). 
These studies indicated that belonging to a large group over the entire 
larval period was not essential and was perhaps even detrimental. 
The group size with highest survivorship to diapause did not exceed 
the average number of eggs per cluster (Stamp, 1981a). 

However, the value of belonging to a large group may fluctuate on 
a daily and even hourly basis as a consequence of larval and parasitoid 


300 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


activity (Stamp, 1982a, b, unpubl. data). For example, newly-hatched 
larvae moved from the oviposition site to the top of the host plant 
stalk, built a communal web, and then began feeding. Without their 
webs and the body spines characteristic of the other instars, first instar 
larvae were particularly vulnerable to parasitoids and predators. One 
value of belonging to a large group was that such a group quickly 
reached the top of the stalk. Newly-hatched larvae from all but the 
first egg cluster deposited on a stalk had a silk trail to follow and a 
communal web to occupy immediately. 

Furthermore, these caterpillars may benefit from membership in 
large groups due to particular defensive mechanisms (e.g. webs, head- 
jerking, unpalatability, aposematic coloration). For example, E. phae- 
ton reared on turtlehead were unpalatable to blue jays (Cyanocitta 
cristata; Bowers, 1980) and, thus, the effect of the aposematic color- 
ation of these larvae may have been enhanced by large group size. 
Also, by head-jerking, second and third instar larvae warded off para- 
sitoids effectively (Stamp, 1982a). If caterpillars in contact with each 
other on the outside of a web were disturbed by a parasitoid, they 
simultaneously head-jerked for several minutes. However, if these 
larvae were not touching each other and were disturbed, only a few 
caterpillars head-jerked and for a shorter period. In the latter case the 
parasitoid continued to search and make contact with caterpillars. 

Since observed survival was highest for moderate-sized groups (that 
is, equivalent to a single egg cluster), why then were many (30%) of 
the clusters in groups? Variation in group size of E. phaeton may be 
the result of two opposing and variable selective pressures. Group 
size of eggs and the ensuing larval aggregations of these checkerspots 
varied tremendously, with one to 10 egg clusters occurring per stalk 
and, thus, about 250 to 2500 newly-hatched caterpillars per web 
(Stamp, 1981a.). By depositing moderate-sized clusters with those of 
other females, E. phaeton may benefit from spreading their eggs, and 
the offspring may benefit during critical larval periods from member- 
ship in large groups. Thus, clumping of egg clusters may enhance 
reproductive success under some circumstances. However, spreading 
eggs may also maximize the probability that some of a female’s eggs 
will survive. 

In conclusion, E. phaeton females were carefully choosing ovipo- 
sition sites and frequently depositing their eggs with those of other 
females. These behaviors suggest a scarcity of particularly attractive 
or high quality stalks. These oviposition behaviors may also indicate 
the value of group membership for larvae during critical periods, such 
as the first instar. 


VOLUME 36, NUMBER 4 301 


ACKNOWLEDGMENTS 


I thank D. H. Morse, R. F. Denno, J. M. Kemper, D. E. Gill and R. S. Fritz for advice 
and comments on the research and manuscript. The research was supported by National 
Science Foundation Grant No. DEB-7907618, Xerces Society, Sigma Xi, University of 
Maryland’s Chapter of Sigma Xi, and the Computer Science Center of the University 
of Maryland. I am grateful to the Conservation and Research Center of the National 
Zoological Park for use of the study area and living accommodations. 


LITERATURE CITED 


BENSON, W. W. 1978. Resource partitioning in passion vine butterflies. Evolution 32: 
493-518. 

BENSON, W. W., K. BROWN, JR. & L. GILBERT. 1975. Coevolution of plants and herbi- 
vores: Passion flower butterflies. Evolution 29:659-680. 

BowERrs, M. D. 1978. Over-wintering behavior in Euphydryas phaeton (Nymphali- 
dae). J. Lepid. Soc. 32:282-288. 

1979. Unpalatability as a defense strategy of checkerspot butterflies with spe- 

cial reference to Euphydryas phaeton (Nymphalidae). Ph.D. Dissertation, Uni- 

versity of Massachusetts, Amherst. 168 pp. 

1980. Unpalatability as a defense strategy of Euphydryas phaeton (Lepidop- 
tera: Nymphalidae). Evolution 34:586—600. 

CHEw, F.S. 1975. Coevolution of pierid butterflies and their cruciferous foodplants. 
I. The relative quality of available resources. Oecologia 20: 117-128. 

1977. Coevolution of pierid butterflies and their cruciferous foodplants. II. 
The distribution of eggs on potential food plants. Evolution 31:568-579. 

EHRLICH, P. R. & L. E. GILBERT. 1973. Population structure and dynamics of the 
tropical butterfly Heliconius ethilla. Biotropica 5:69-82. 

GILBERT, L. E. 1975. Ecological consequences of a coevolved mutualism between 
butterflies and plants. Pp. 210-240 in L. E. Gilbert and P. H. Raven (eds.). Co- 
evolution of animals and plants. University of Texas, Austin. 

LABINE, P. A. 1968. The population biology of the butterfly, Euphydryas editha. VIII. 
Oviposition and its relation to patterns of oviposition in other butterflies. Evolution 
22:799-805. 

RAUSHER, M. D. 1979. Egg recognition: Its advantage to a butterfly. Anim. Beh. 27: 
1034-1040. 

1980. Host abundance, juvenile survival, and oviposition preference in Battus 
philenor. Evolution 34:342-355. 

ROTHSCHILD, M. & L. M. SCHOONHOVEN. 1977. Assessment of egg load by Pieris 
brassicae (Lepidoptera: Pieridae). Nature (Lond.) 266:352-355. 

SOUTHWOOD, T. R. FE. 1978. Ecological methods. 2nd ed. Chapman & Hall, London. 
524 pp. 

STAMP, N. E. 1979. New oviposition plant for Euphydryas phaeton (Nymphalidae). 
J. Lepid. Soc. 33:203-204. 

1980. Egg deposition patterns in butterflies: Why do some species cluster 

their eggs rather than deposit them singly? Amer. Natur. 115:367-380. 

198la. Effect of group size on parasitism in a natural population of the Bal- 

timore checkerspot Euphydryas phaeton. Oecologia 49:201—206. 

1981b. Parasitism of single and multiple egg clusters of Euphydryas phaeton 

(Nymphalidae). J. N.Y. Entomol. Soc. 89:89-97. 

1982a. Behavioral interactions of parasitoids and Baltimore checkerspot cat- 

erpillars (Euphydryas phaeton). Environ. Entomol. 11:100-104. 

1982b. Searching behaviour of parasitoids for web-making caterpillars: A test 

of optimal searching theory. J. Anim. Ecol. 52:387-395. 

1982c. Aggregation behavior in Baltimore checkerspot caterpillars Euphydryas 

phaeton (Nymphalidae). J. Lepid. Soc. 36:31-41. 


302 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


WHITHAM, T.G. 1978. Habitat selection by Pemphigus aphids in response to resource 
limitation and competition. Ecology 59:1164—-1176. 

WIKLUND, C. 1974. The concept of oligophagy and the natural habitats and host plants 
of Papilio machaon in Fennoscandia. Entomol. Scand.. 5:151—160. 

1975. The evolutionary relationship between adult oviposition preferences 
and larval host plant range in Papilio machaon L. Oecologia 18:185-197. 

WILLIAMS, E. H. 1981. Thermal influences on oviposition in the montane butterfly 
Euphydryas gillettii. Oecologia 50:342-346. 


Journal of the Lepidopterists’ Society 
36(4), 1982, 303 


GENERAL NOTES 


FIELD OBSERVATIONS OF DIVERGENT RESTING BEHAVIOR AMONG 
HICKORY FEEDING CATOCALA LARVAE (NOCTUIDAE) 


Catocala are known for their sympatric diversity, with at least 35 species present at 
almost any forested locality in southern New England (see Sargent, 1976, Legion of 
night: The underwing moths, Univ. Massachusetts Press, Amherst, MA; also personal 
observation). Of these, about one third feed on Juglandaceae (hickories and walnuts). 
The precise foodplant preferences of these Juglandaceae feeders will be the topic of 
another paper (Schweitzer, in prep.) and are also being investigated by L. F. Gall. 

Sargent (1976, op. cit.; 1977, J. Lepid. Soc. 31:1—-16) suggested that food may not be an 
important limiting factor in Catocala evolution. “Competition” for predator avoidance 
might be more important in Catocala ecology. Such competition might be manifest in 
different larval resting behaviors, which might serve to inhibit effective search image 
formation by predators, especially birds. Sargent (1976, op. cit.) reports laboratory obser- 
vations of instar specific resting behaviors in Catocala dejecta Strecker and C. retecta 
Grote which were reared in plastic containers on Carya ovata. Although similar in ap- 
pearance the two species grow at very different rates and so avoid having similar size 
and behavior simultaneously. 

Little is known about field resting habits of most Catocala species. From my own 
observations it seems that late instar C. epione (Drury) and C. c. consors (J. E. Smith) 
larvae almost invariably rest on branches or small trunks. Late instars of Catocala 
habilis larvae are quite flattened and can be found resting under bark shags. Most 
sleeved C. palaeogama and C. dejecta last instar larvae rested exposed on the branches. 
Most other species typically hid in folds of the cloth or in debris. 

The resting habits of larvae collected on 25 and 26 May 1980, West Rock, New Haven, 
Connecticut, were noted and are presented in Table 1. Data for those larvae observed 
at rest were analyzed by chi-square tests. Catocala epione differs from the other species, 
all of which appear to have similar resting behavior (x? = 10.6, P < 0.005, with Yates 
correction). Of course, resting habits may diverge in later instars, and a larger sample 
size might have revealed more differences. 

These limited observations are apparently the first demonstration of differences in 
field resting behavior of the sort described by Sargent and may be the first published 
documentation of natural foodplants for these species. The data in Table 1 are from 
larvae collected on four stunted trees at the edge of trap rock outcrops. All trees had 
trunk diameters of less than 15 cm and had some bark exfoliations on the trunk. All 
branches were carefully searched and then beaten with a baseball bat. All but two of 
the 25 larvae were found by the visual search. 


DALE F. SCHWEITZER, Curatorial Affiliate, Entomology Section, Peabody Museum, 
P.O. Box 6666, Yale University, New Haven, Connecticut 06511. 


TABLE 1. Resting behavior of Catocala larvae collected at West Rock, New Haven, 
Connecticut on 25 and 26 May 1980 on Carya ovata. C. epione were in ultimate (1) and 
penultimate (5) instars. Others were 2-3 cm long and dark colored, and were probably 
in the third instar. 


Numbers observed resting on 


Beaten Foliage! Twig Trunk Crawling 
C. epione 1 0 5 0 0 
C. palaeogama 1 9 1 0) 1 
C. residua 0 4 0 0 0 
C. retecta 0 a il 0 0 


' Midrib on underside of leaflets. 


Journal of the Lepidopterists’ Society 
36(4), 1982, 304-307 


A LATE-SEASON EMERGENCE OF CALLOPHRYS (SANDIA) 
MACFARLANDI (LYCAENIDAE) 


Callophrys (Sandia) macfarlandi Ehrlich & Clench is extremely restricted in larval 
foodplant utilization, although it is widely distributed and often common in much of 
New Mexico east of the Continental Divide (Holland, 1974, J. Lepid. Soc. 28:38-52). 
It appears to be an obligatory feeder on Nolina texana Wats. (Liliaceae). In the small 
portion of New Mexico which is south and west of Truth or Consequences, N. texana 
is replaced by N. microcarpa Wats., seemingly without intermediate forms. The two 
species of Nolina never occur together in New Mexico, although they come within 30 


Precipitation (cm) 


BS) '60 OS 0 


Fic. 1. July rainfall at Sunspot, NM, for 1955-1974. 


VOLUME 36, NUMBER 4 305 


Fic. 2. 9 (upper) and 6 (lower) Callophrys (Sandia) macfarlandi, ventral surfaces, 
19-VIII-74, Grapevine Can., Sacramento Mts., Otero Co., NM, 6000’ (1640 m). 


mi. (50 km) of contact in some places. Callophrys macfarlandi has never been observed 
in association with N. microcarpa. On the other hand, whenever one locates a stand 
of N. texana, no matter how isolated from other stands, one can nearly always find 
macfarlandi. It has not yet been determined whether macfarlandi will accept micro- 
carpa in captivity when no choice is offered. The most obvious distinguishing feature 
between the two species of Nolina is the height of the inflorescence. In texana, the 
inflorescence is about the same height as the leaf, while in microcarpa the inflores- 
cence is approximately twice the leaf height. This gives microcarpa some resemblance 
to Yucca, a genus closely related to Nolina. 

Callophrys macfarlandi larval feeding is restricted not only to N. texana but, in 
particular, to the bloom of the plant. Nolina texana normally blooms in April or early 
May, depending on the location. Fresh macfarlandi have been captured from 15 Feb- 
ruary to 29 June, however, with no clear pattern of discrete broods. This seasonal 
distribution raises some difficult questions. First, it is uncertain how many generations 
of macfarlandi occur in a given year. Blooms stay fresh long enough and the macfar- 
landi larval stage is brief enough that two complete generations may be possible. 
However, there is definitely no bloom available under any circumstances in June. This 
seemed to mean the late May and June emergers are either a “suicide brood” or the 
species is able to pass the fall and winter as ova. One would normally expect natural 
selection to act very swiftly against genotypes with a “suicide brood” proclivity. This 
viewpoint would favor the fall-winter, ova-diapause hypothesis. The problem with that 


306 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Precipitation (cm) 


a5 "60 OS 20 


Fic. 3. June rainfall at Sunspot, NM, for 1955-1974. 


hypothesis is it is incompatible with the February flight, unless mag onan can dia- 
pause either as ova or pupa. 

A purely speculative explanation for these observations may be found in a series of 
events which took place in 1974. This year was marked by an exceptionally wet July 
around Alamagordo, Otero Co., NM. Rainfall records were kept for.a 20-year period 
from August 1954, to July 1974, at Sunspot, NM, high in the Sacramento Mts. near 
Alamagordo (Demastus, 1976, A twenty-year summary of Sacramento peak weather— 
August 1954 through July 1974, Sacramento Peak Observatory, Air Force Geophysics 
Laboratory, Hanscom AFB, Massachusetts 01731, AFGL-TR-76-0096, Special Report 
No. 196). July rainfall at Sunspot is graphed in Fig. 1 for these years. It may be seen 


VOLUME 36, NUMBER 4 307 


that July 1974 was significantly wetter than any other July between 1964 and 1974. 
(These eleven seasons include the ones in which I had been active in New Mexico 
and consequently, for which I have overlapping data both on rainfall and Lepidoptera.) 

In August 1974 around the Sunspot-Alamagordo area, the Nolina did a very unex- 
pected thing, apparently in response to the extreme wetness of July: Many of the plants 
bloomed a second time. The August inflorescence was quite dwarfed compared to the 
normal bloom; its height was perhaps one fourth of the height of the spring inflores- 
cence. 

On 19 August 1974, I checked a summit in the Sacramento Mts. foothills about 5 mi. 
(8 km) SE of Sunspot. This particular summit has often produced interesting hilltopping 
specimens, although macfarlandi does not usually hilltop. In this case however, I was 
rewarded by the capture of a fresh female and an extremely worn male (Fig. 2). Despite 
the tattered condition of the male, it seems unlikely this small lycaenid could have 
been on the wing since June. 

If we again refer to Fig. 1, we see that the July 1955 and 1960 precipitation was also 
great enough at Sunspot to have caused the Nolina to bloom again in August, had 
anyone been there to look. I, thus, suggest that the macfarlandi which emerge in June 
may be favored by virtue of their progeny being able to utilize the August flowering 
of their foodplant if very heavy July rains intervene. If this strategy is successful three 
years out of twenty, as our limited data here indicates, the selection pressure against 
June emergence would, at least, be considerably mitigated. Normally, June is far drier 
than July in the Alamagordo area; however, highly unusual rains in June 1966, could 
conceivably have triggered a second blooming of Nolina that year also (see Fig. 3). If 
this indeed occurred, the odds for the June emergence successfully completing a life 
cycle without diapausing as ova improve to four in twenty. 

On the other hand, it is fully possible that the “suicide brood” concept is actually 
correct. For instance Shapiro (1967, J. Res. Lepid. 6:181—-183) has demonstrated that 
in Pennsylvania Colias and Pieris are subject to suicide eclosions just before the onset 
of lethally cold weather. Also, the presence of a female macfarlandi exhibiting hill- 
topping (mate-seeking) behavior implies the number of available males at that time 
and place must have been very low. Whether by accident or adaptation, however, the 
unseasonal concurrence of Nolina bloom and macfarlandi eclosion in August 1974 is 
a most interesting example of desert survival. 

The American Museum of Natural History has macfarlandi specimens of both sexes 
taken by F. H. Rindge at Sitting Bull Falls near Carlsbad, Eddy Co., NM, on 27 and 
29 July 1964. I am not personally familiar with that site and feel unqualified to speculate 
on a relationship between this record and my own observations. 


ACKNOWLEDGMENTS 


J. McCaffrey formerly of Sunspot, NM, and now at New Mexico State University, 
Las Cruces, succeeded in tracking down the obscure Air Force precipitation report 
which made a cohesive presentation of these items possible. 


RICHARD HOLLAND, 1625 Roma NE, Albuquerque, New Mexico 87106. 


Journal of the Lepidopterists’ Society 
36(4), 1982, 308-309 


THE CORRECT PLACEMENT OF EVERES ‘HERRIT 


Since its description in 1901, the taxonomic placement of herrii (Grinnell) has been 
in question. After being variously designated as a subspecies of Everes comyntas (Go- 
dart) and Everes amyntula (Boisduval), what are believed to be the type specimens 
have now been examined genitalically and herrii’s alignment with amyntula is finally 
decided. 

Grinnell (1901, Can. Entomol. 33:192) described Lycaena amyntula var. herrii from 
“two males and two females” which were collected by Poling in July 1899 and Sep- 
tember 1900. The original description of the male states that it differs from typical 
amyntula “... in having a black margin about 1 mm wide; whereas, there is none in 
typical amyntula, or, if any, a very slight trace. On the underside the markings are 
much more heavy. The male of this variety is also much smaller than the male of typical 
amyntula ....’ And the description of the female states, “... differs from typical 
amyntula by the replacement of the dark area of the primaries by a narrow black band 
about 1 mm wide, and on the secondaries by only two red crescents instead of five as 
in typical amyntula.” 

Then, Bethune-Baker (1913, Ent. News XXIV:97), while not questioning the place- 
ment of herrii within the genus, argued that the differences between amyntula and 
herrii were flimsy and that if Grinnell “... had had the advantage of having the type 
of amyntula before him that he would not have described the form.” 

The actual comyntas/amyntula debate was begun by Bares and McDunnough (1916, 
Cont. Nat. Hist. N.A. III:109), who subsequently state that after examining a large series 
of Everes from southeastern Arizona, “... we should be inclined to refer herri Grinnell 
to comyntas rather than amyntula ....” They state that this decision is based in part 
on the original description, which mentions the “. . . broader black border on the upper 
side and the better defined and larger spots on the under side ...”’; in part on the 
presence of “... red lunules near the anal angle on the secondaries above .. .” in many 
of the specimens, a feature closely associated with comyntas; and their decision is also 
based in part on the fact that the dates of capture point to a distinct double-broodedness. 
This multiple- versus single-brooded aspect had been suggested by Bethune-Baker 
(op. cit.), who, however, alluded to a partial second brood in amyntula in the southern 
end of its range. 

Aside from the opinions of these early researchers as to the placement of herrii, 
another issue complicated matters. Were, in fact, amyntula and comyntas actually 
distinct? Because, if not, the herrii point was moot. Various writers, including Holland, 
Klots, Brown et al., dos Passos, and Howe, follow the separation of American Everes 
into the two taxa, while a smaller number, including Ehrlich, have lumped the two 
under comyntas. So, in several ways, the placement of herrii has been questionable. 

However, what was not known until recently was the fact that both a comyntas and 
an amyntula “type” occur in southeastern Arizona, specifically, in Cochise County and 
environs. So, when Downey and Christenson (1970, Proc. No. Cen. Branch—E.S.A. 
25(2):89) reexamined the male genitalia of the American Everes and reaffirmed the fact 
that there are indeed “... 2 discrete morphological types (species) ...,” the genitalic 
identity of the “types” of herrii needed to be disclosed. 

At this point Julian Donahue of the County Museum in Los Angeles (LACM) proved 
to be most helpful. He located the four specimens which seemed to match the four 
mentioned by Grinnell (op. cit.), “. . . with notes by Comstock that they are probably the 
true types.” Oddly, these four specimens were found to be all males. Donahue then 
dissected the two specimens taken in July 1899. The other two were sent to me. (The 
lower label on these two specimens reads “Lyc. amyntula v. herrii,’ while the upper 
label says “So. Ariz. Sept 1900. Poling.”) All four were found to be amyntula. The 
smaller of the two males taken September 1900 is hereby designated as the lectotype. 

Bethune-Baker (op. cit.) stated that the original four specimens were taken in Cochise 
County. To get more specific than that requires some conjecture. All known males of 
Everes from the Chiricahua and Peloncillo mountains have proven to be comyntas. A 


VOLUME 36, NUMBER 4 309 


female in the LACM collection taken by V. W. Owen in the Chiricahua Mtns., 9 May 
1910, has considerable blue dorsal scaling and does appear to be an amyntula. Barnes 
and McDunnough (op. cit.) based their work on a large series from the “... Huachuca 
Mtns. and other mountain chains of S.E. Ariz.” Since the publication of their work 
some 65 years ago, only a handful of Everes has been taken in the Huachuca Mtns., 
all of which have been amyntula. Also, amyntula has been confirmed from the Pata- 
gonia Mtns., just west of the Huachucas, but in Santa Cruz County. In light of the 
apparent absence of comyntas from the Huachuca Mtns., the mention of Barnes and 
McDunnough (op. cit.) of their ‘large’ series from that area, and the confirmed occurrence 
of amyntula in that area, it is here suggested that the Huachuca Mtns. are the probable 


type locality of herrii. 


RICHARD A. BAILOWITZ, 1750-B Xavier Way, Nogales, Arizona 85621. 


Journal of the Lepidopterists’ Society 
36(4), 1982, 310-314 


NATURAL HISTORY OF HYPNA CLYTEMNESTRA CR. 
(NYMPHALIDAE) IN COSTA RICA 


With the exception of several old reports (e.g., Muller, 1886, Zool. Jahrb. Zeitschr. 
Syst. Geogr. Biologia der Tiere, Jena 1:417-678; Rober, 1914, In Seitz, Macrolepidop- 
tera of the World, Vol. 5, Stuttgart, A. Kernan Verlag, 516 pp.) and a few recent studies 
(e.g., Muyshondt, 1973a, J. Lepid. Soc. 27:294-302; Muyshondt, 1974, J. Lepid. Soc. 
28:81-89; Muyshondt, 1975, J. Lepid. Soc. 29:168-176), little has been published on 
the larval host plants and early stages of the Neotropical Charaxinae or Charaxidae 
(Nymphalidae). Emphasis is given to this fact by the statement of Rober (op. cit.) that 
only the pupa stage is known for the three species of Hypna. It is generally known, 
however, that the closely allied genus Anaea exploits a variety of larval host plants in 
different families, including the Lauraceae, Flacourtiaceae, and Euphorbiaceae (Muy- 
shondt, 1973a, 1974, 1975, op. cit.; Young, 198la, Acta Oecologia 2: 17-30), and it would 
therefore, not be surprising to discover a similar feeding pattern for Hypna. In this 
note I report the description of larval and pupal stages of Hypna clytemnestra Cramer 
from one locality in Costa Rica, and also report the larval host plant to be Croton sp. 
(Euphorbiaceae). Both Rober (op. cit.) and Riley (1975, A field guide to the butterflies 
of the West Indies, London, Quadrangle, 224 pp.) confirm the lack of data on larval 
host plants of the genus as well as information on the early stages. 

In Costa Rica, H. clytemnestra is very rare on the eastern slopes of the Cordillera 
Central (based on observations by Young, 1968-1982). I have captured a few adults at 
“Finca La Tigra,” near La Virgen (10°23’N, 84°07’W; 220 m elev.), Heredia Province, 
during the short dry season (February—March). On 19 February 1981 a single “Anaea- 
type” caterpillar was discovered on a 4-meter tall Croton sapling (Fig. 1) at the edge 
of a steep forest ravine at Turrialba (9°54’N, 83°41'W; 602 m elev.), Cartago Province. 
As with the La Tigra site, this region is classified as Premontane Tropical Wet Forest. 
This caterpillar was subsequently reared by confining it with fresh host plant cuttings 
in a large clear-plastic bag. The single host plant individual was searched for additional 
caterpillars, but none were found. At the time of its discovery, the caterpillar was in 
the third instar. The ant, Zacryptocerus scutulatus (F. Smith), was abundant on the 
leaves and stems of the plant but did not interact with the Hypna caterpillar. 

The single individual of Croton sp. with the caterpillar was the only one found within 
a 400-meter strip of the forest perimeter checked. 

Following eclosion, the butterfly was identified as Hypna clytemnestra and probably 
the subspecies clytemnestra Cramer, given discussion of geographical distribution (e.g., 
Comstock, 1961, Butterflies of the American tropics, the Genus Anaea, Lepidoptera, 
Nymphalidae, New York, Amer. Mus. Nat. Hist., 214 pp.). In captivity the larva molted 
twice, suggesting that it was in the third instar at the time of discovery, since related 
forms typically have five instars (Muyshondt, 1973a, 1974, 1975, op. cit.; Young, 1981a, 
op. cit.). The third instar grew from 15 mm to 23 mm in five days and remained mottled 
in shades of green and brown. The head is studded with tiny whitish tubercles and 
one pair of stubby horns on the apex of the epicrania (Fig. 2). Alternating thoracic and 
abdominal segments bear bulbous tubercles of varying size and studded with tiny 
tubercles (Fig. 2). The spiracles are black. Third and fourth instars construct a silk- 
encased perch from the exposed midrib of a mature leaf of the food plant (Fig. 2). The 
larva rests and feeds from this perch (Fig. 2). In the fifth instar, which attains a length 
of 41 mm by the time of pupation, the thoracic and abdominal tubercles are reduced 
in size, but now each one bears a thick black spine (Fig. 2). The head is laterally 
flattened, with many tubercles (Fig. 2). In this instar the larva, in captivity, rests on 
the ventral side of the leaf. When disturbed or walking, the larva typically wobbles 
from side to side, walking forward a bit, and then backwards a bit, before moving ahead. 
The “wobbling” motion is also exhibited when the larva is reinforcing its silken perch 
with additional threads of silk. The head capsules of the fourth and fifth instars are 
shown in Fig. 3. 

Pupation involves the larva assuming the “J” position, in this instance from the stem 


VOLUME 36, NUMBER 4 311 


Fic. 1. The sapling-size individual of Croton sp. (Euphorbiaceae) where a single 
third instar larva of Hypna clytemnestra was discovered. The plant in question is 
shown here immediately above the butterfly net. Note the large heart-shaped leaves 
of the sapling. See text for locality and habitat information. 


of a leaf. As a prepupa the larva remains tightly curled, and the pupa is initially uniform 
green. Within twenty-four hours, however, the pupa (Fig. 4) turns dark green and 
develops a patchwork of silvery pubescence, particularly on the wing pads (Fig. 4). 
The pupa measures 20 mm long by 13 mm at its widest girth and laterally. The head 
is slightly forked, spiracles are yellow, and the cremaster red. At the posterior edge of 


SLY, JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


in resting and feeding positions on the midrib perch at the tip of a leaf. Bottom, left 
to right: fifth instar. 


Fic. 3. Frontal view of head capsules of fourth (rt.) and fifth instars of the larvae of 
Hypna clytemnestra. 


VOLUME 36, NUMBER 4 313 


Fic. 4. Pupa, eclosion, and adult Hypna clytemnestra. Top, left to right: pupa 
ventral and lateral views respectively, and newly-eclosed adult resting adjacent to 
empty pupa shell. Bottom, left to right: dorsal and ventral views of the single reared 
adult. 


the fifth abdominal segment there is a prominent ridge, marked by a thin line of gold 
color. On the day of eclosion the adult butterfly is seen clearly through the cuticle. 
The pupa stage lasted 16 days at the Milwaukee Public Museum room temperature 
(about 23°C at 3 PM). Eclosion (Fig. 4) is rapid, lasting about three hours. The adult 
obtained was a male with forewing length of 42 mm (Fig. 4). 

This report constitutes the first description of the larval and pupal stages of H. cly- 
temnestra and a larval food plant record. The Neotropical Charaxinae exploit a variety 
of plant families as larval food plants (Ehrlich & Raven, 1965, Evolution 18:586—608; 
Muyshondt, 1973a, 1974, 1975, op. cit.; Young, 1981a, op. cit.) and Muyshondt (1975, 
op. cit.) reported Anaea (Memphis) pithyusa R. Felder feeding on Croton in El] Sal- 
vador. Other genera or subgenera are associated with other plant families. To this list 
we add that Hypna, although very distinct from Memphis in terms of the appearance 
and behavior of the early stages (larva and pupa) as well as adult characteristics, also 
exploits Croton. Furthermore, some genera or subgenera within the Charaxinae or 
Charaxidae may be associated with more than a single plant family. Young (198 1a, op. 
cit.) found Anaea (Memphis) morvus feeding on Nectandra sp. (Lauraceae) in Costa 
Rica, while A. (M.) pithyusa feeds on Croton (Euphorbiaceae) in El Salvador (Muy- 
shondt, 1975, op. cit.). Geographical changes in larval food plant associations at the 
genus or subgenus levels, or changes in such associations for different species may be 
operative, although further studies are required. Such data points to the difficulty in 
using food plant data in the determination of evolutionary lineages at levels lower than 
families or subfamilies in the Lepidoptera. Contemporary selection pressures may have 


314 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


obliterated or changed the original food plant association (Janzen, 1980, Evolution 34: 
611-612). 

The distinctive dorsal tubercles of the larva of Hypna, and its characteristic “wob- 
bling’ behavior, represent two other phenotypic traits very different from most other 
Charaxinae. Midrib-perching is shared with other subgenera such as Zaretis and Con- 
sul (see Muyshondt, 1973a, 1974, op. cit.; Young, 1981a, op. cit.). Although Hypna is 
widespread in South America (Comstock, op. cit.) it is predicted to be associated with 
Croton through its range. There are no published records of H. clytemnestra from 
Costa Rica, although the form clytemnestra clytemnestra is known from Nicaragua and 
Panama, and also ranges from Colombia to Brazil (Comstock, op. cit.). As with most 
other tree or vine-exploiting nymphalids, and particularly the Nymphalinae and Cha- 
raxinae or Charaxidae (e.g., Young, 1981b, J. Lepid. Soc. 35: 155-157), this species occurs 
primarily in tropical wet forests. 

Muyshondt (1973b, J. New York Entomol. Soc. 81:164—174) reports the larval food 
plant of Catonephele numilia esite Felder in El Salvador to be Alchornea latifolia 
Swartz in the Euphorbiaceae, and places this species within the Catonephelinae. Such 
discoveries indicate the joint exploitation of the same plant family by different evo- 
lutionary lineages within the Nymphalidae. As discussed by Muyshondt (1975, op. cit.), 
Rydon (1971, Entomol. Rec. J. Var. 83:219-388) places the subgenera Hypna, Anaea, 
Polygrapha, Consul, and Memphis in the subfamily Anaeinae as full genera. As men- 
tioned above in terms of geographical and ecological determinants of larval food plant 
associations in Neotropical nymphalids, the euphorbiaceous-feeding habit of the larval 
stages of Hypna and Memphis probably cannot be used to support the view of a close 
evolutionary association between these two genera or subgenera (the latter, adopting 
Comstock’s 1961 classification). Furthermore, the larva of Hypna is quite distinct both 
morphologically and behaviorally from that of Memphis (see Muyshondt, 1975, op. cit.). 
The pupa stage of Hypna is closer in general appearance to that of Consul (Muyshondt, 
1974, op. cit.) than it is to that of Memphis, yet Consul is a piperaceous-feeder (Muy- 
shondt, 1974, op. cit.). 


ACKNOWLEDGMENTS 


This research was funded through a grant from The American Cocoa Research In- 
stitute. I thank Dr. Gary S. Hartshorn, Tropical Science Center, Costa Rica, for iden- 
tifying the food plant, and Ms. Susan Sullivan Borkin for assistance with finding dis- 
tributional information on the butterfly. I thank Dr. Robert Murray for assistance in 
preparing Fig. 3. Special thanks to Jorge W. Coto (Costa Rica) and Leo Johnson (Mil- 
waukee Public Museum) for photographic assistance. The single reared specimen from 
this study is deposited, along with larval head capsules and pupa shell, in the Lepi- 
doptera collections of the Milwaukee Public Museum. This paper is dedicated to Al- 
berto Muyshondt, a pioneer in the study of life cycles and natural history of the but- 
terflies of El Salvador. 


ALLEN M. YOUNG, Invertebrate Zoology Section, Milwaukee Public Museum, Mil- 
waukee, Wisconsin 53233. 


Journal of the Lepidopterists’ Society 
36(4), 1982, 315-319 


INDEX TO VOLUME 36 


(New names in boldface) 


Acsala anomala, 218 Atrophenura sycorax, 56 
Adler, P. H., 161 A. varuna, 56 
Aechmea bracteata, 70 Atrytonopsis deva, 157 
Agonopterix alstroemeriana, 160 A. python, 157 

A. clemensella (host plants), 160 Audre domina, 71 
Agraulis vanillae, 246 Bailowitz, R. A., 308 
Agriades rustica rustica, 158 Bembecia bestianaeli, 120 
Agrocercops striginifinitella (?), 163 B. chrysidiformis, 120 
Aiello, A., 65 B. dancaudani, 120 
Ailanthus altissima, 198 B. hannemanni, 120 
Alypia octomaculata, 237 B. ili, 120 
Amata marjana, 188 B. polistiformis, 120 

A. phegea, 185 B. reisseri, 120 

A. ragazzii, 185 B. sanguinolenta, 120 


Amathusia spp., 56 Berenbaum, M., 160 
Ameiva ameiva, 149 Bianco, P., 185 
Amplypterus gannascus, 155 Biston betularia cognataria, 163 
Amyris elemifera, 254 Blepharomastix ranalis, 163 
Anacamptodes ephyraria, 163 Boloria astarte distincta, 223 
Anaea moretta, 202 Bolte, K. B., 83 
Anagoga occiduaria, 163 Bomolocha baltimoralis, 164 
Ananas cosmosus, 70 Book Reviews, 216, 240 
Anania funebris glomeralis, 163 Brenthia dendronympha, 120 
Anartia jatrophae, 246 . B. spintheritis, 120 
Anavitrinella pampinaria, 163 Brephidium pseudofea, 247 
Ancylis metamelana, 163 : Brower, A. E., 159 
Anguria tabascensis, 179 Bryant, R. S:,237 
Antepione thisoaria, 163 Bucculatrix ulmicola, 145 
Anthocharis cardamines, 132 B. ulmifoliae group, 145 
Antigonum leptopus, 69 Buellia cf. spuria, 225 
Antirrhinum sp., 33 Bullini, L., 185 
Aphelandra depeana, 67 Callicore hydaspes, 148 
Aphrissa statira, 248 Callophrys macfarlandi, 304 
Appias drusilla, 249 Callosamia angulifera, 237 
A. indra, 56 C. promethea, 154, 192, 208, 237 
A. lyncida, 56 Caloptilia cornusella, 163 
A. nero, 56 Cannell, P. F., 264 
Araucaria cunninghamii, 269 Capparis pittieri, 229 
Arawacus aetolus lincoides, 66 Carter, N. G., 240 
Archaeoattacus edwardsii, 112 Carya cordiformis, 259 
Archonias tereas, 203 C. glabra, 257, 258 
Argyria nivalis, 163 C. illinoiensis, 261 
Arnaud, P. H., 62 C. ovalis, 258 
Artogeia rapae, 133, 266 (Pieris) C. ovata, 47, 257, 258 
Asbolis capucinus, 249 C. tomentosa, 258 
Ascia monuste, 249 Catocala andromache, 159 
Aster spp., 121 C. benjamini, 159 
Asterocampa celtis, 235 C. blandula, 22 
A. clyton, 234 C. crataegi, 18 
Astraptes crana, 236 C. dejecta, 262, 303 


A. fulgerator, 236 C. epione, 258, 303 


316 


C. flebilis, 262 
C. habilis, 258, 303 
C. judith, 256 
. maestosa, 258 
mira, 21 
. neogama, 42, 258 
. obscura, 259 
palaeogama, 257, 303 
pretiosa, 18, 256; Status Change, 23 
. residua, 257, 303 
. retecta, 42, 259, 303 
robinsonii, 256 
. serena, 261 
. subnata, 259 
. texarkana, 22 
. vidua, 258 
Celtis laeviga, 234 
Cenopis reticulatana, 163 
Cepora nadina, 56 
Cepphis armataria, 163 
Ceratomia amyntor, 237 
Cercyonis pegala, 265 
Chamaesphecia balcanica, 120 
C. corsica, 120 
C. meriaeformis, 120 
C. monspeliensis, 120 
C. rangnovi, 120 
C. tengyraeformis, 120 
Charaxes bernardus, 56 
Chelone glabra, 31 
Chlorochlamys chloroleucaria, 163 
Chlosyne harrisii, 265 
Cianchi, R., 185 
Cilek, J. E., 227 
Citheronia regalis, 76, 237 
C. splendens, 80 
Clemensia albata, 222 
Clostera albosigma, 164 
Colias crocea, 133 
C. eurytheme, 266 
C. interior, 266 
Cooper, W. J., 207 
Cordia sebestena, 67 
Cornutia grandifolia, 68 
Coryphista meadii, 163 
Courtney, S., 132 
Crambus turbatellus, 163 
Crataegus uniflora, 20 
Croton bilbergianus, 72 
C. niveus, 71 
Cupha erymanthis, 56 
Cyanocitta cristata, 196 
Cyanophrys herodotus, 68 
Cyclophora myrtaria, 163 
C. packardi, 163 
Cydista sp., 68, 69 
Cyrestis nivea, 56 


VAIQGAMIAQASGVAGgaaG 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Danaus aspasia, 56 

D. gilippus, 245 

D. plexippus, 245, 265 
Datura arborea, 67 
De Jong, R., 279 
Delias descombesi, 59 
De Matthaeis, E., 185 
Desmia funeralis, 163 
Desmodium axillare, 70 
De Vries, P. J., 229 
Dioclea guinensis, 69 
Dione juno, 137, 245 
Diospyros virginiana, 76 
Dipchasphecia roseiventris, 119 
Discophora timora, 58 
Dryas julia, 137, 245 
Duckworth, W. D., 119 
Ductispina turcmena, 119 
Durden, C. J.—Barton Creek Butterflies, 

1 

Dynamine artemisia, 148 
Dyspteris abortivaria, 163 
Dysstroma hersiliata, 164 

D. h. “‘mirandata’’, 83 
Ectatomma ruidum, 66 

E. tuberculatum, 66, 71 
Ectropis crepuscularia, 164 
Ehrlich, A. H., 148 
Ehrlich, P. R., 148 
Enargia decolor, 164 
Epargyreum zestos, 250 
Erebia epipsodea, 203 

E. magdalena mackinleyensis, 223 
Erynnis horatius, 227 

E. zarucco, 250 
Euchlaena irraria, 164 
Eudeilinea herminiata, 163 
Eueides aliphera, 137 

E. isabella, 137 

E. lineata, 136 

E. lybia, 137 

E. tales, 143 

E. vibilia, 136 
Eunica margarita, 149 

E. tatila, 246 
Eupackardia calleta, 207, 237 
Euphydryas aurinia, 299 

E. chalcedona, 151 

E. editha, 298 

E. gillettii, 299 

E. phaeton, 31, 290 
Euphyia unangulata intermediata, 164 
Eupithecia sp., 164 
Euploea camaralzeman, 56 

E. diocletianis, 56 

E. mulciber, 56 


VOLUME 36, NUMBER 4 317 


Euptychia areolata, 245 Immature Stages (descriptive): 
E. hermes sosybia, 235 Ova, 19, 139, 140 
Eurema daira, 249 Larva, 19, 139, 142, 222, 310 
Eusarca confusaria, 164 Pupa, 139, 142, 211 
Everes comyntas var. herrii, 308 Inga pezizifera, 71 
Faunis gracilis, 56 Tridopsis larvaria, 164 
Ferge, L. A., 233 Isatis tinctoria, 132 
Ferguson, D. C., 218 Israel, M. L., 227, 234 
Flynn, D. J., 157 Itame abruptata, 159 
Forestiero, S., 185 I. argillacearia, 164 
Franclemont, J. G., 218 I. pustularia, 164 
Freeman, H. A., 236 Juglans cinerea, 248 
Gandaca harina, 56 J. nigra, 248 
Gelis obesus, 226 Kane, S., 200 
Gilbert, L. E., 178 Kohleria tubiflora, 70 
Glaucopsyche lygdamus, 266 Lafontaine, J. D., 218 
Gliricidia sp., 69 Lantana camara, 67, 68 
Gluphisa septentrionis, 164 Larsen, T. B., 238 
Gonepteryx rhamni, 133 Lecidia armeniaca, 225 
Graphium antiphates, 56 L. fuscocinerea, 225 
G. evemon, 56 Lenczewski, B., 241 
G. macareus, 56 Leptosa nina, 56 
Grapholita eclipsana, 163 Leptotes cassius, 248 
Gupta, M. L., 112 Leston, D., 241 
Haematopis grataria, 164 Leucospilapteryx venustella, 163 
Hamadryas ferentina, 202 Limenitis archippus, 265 
Heliconia latispatha, 69 L. arthemis, 265 
H. wagneriana, 69 L. astyanax, 235 
Heliconius charitonia, 178, 245 Lippia sp., 121 
H. doris, 182 Liquidambar styraciflua, 76 
H. erato, 182 : Lithochodia carneola, 165 
H. hewitsoni, 141 Lobophora nivigerata, 164 
H. ismenius, 182 Lomographa semiclarata, 164 
H. sara, 143 Longino, J. T., 136 
Heliomata cycladata, 164 Lycaeides argyrognomon nabokovi, 233 
H. infulata, 164 Lycaena epixanthe, 266 
Hemiargus ceraunus, 248 L. phleaus americana, 266 
H. thomasi, 248 Maddox, G. D., 264 
Hemileuca maia, 237 Malacosoma americanum, 39 
Heppner, J. B., 87, 119 Mallet, J. L. B., 136 
Heterophleps triguttaria, 164 Mangifera indica, 67, 68 
Hibiscus rosa-sinensis, 68 Manley, T. R., 60 
H. tubiflorus, 70 Marpesia chiron, 149 
Highfill, F. W., 207 M. petreus, 149, 246 
Hodges, R. W., 216 Mascagnia hippocratioides, 68 
Holland, R., 304 Mather, B., 159 
Homochlodes fritillaria, 164 Mechanitis isthmia, 55, 66 
Homogyna porphyractis, 120 M. lysimia macrinus, 66 
Hyalophora cecropia, 60, 154, 192, 207 Megisto cymela cymela, 153 
Hydrelia lucata, 164 M.c. viola, 153 
Hydria prunivorata, 164 Melanitis leda, 56 
Hydriomena perfracta, 164 Melanocyma faunula, 56 
Hypena humuli, 165 Melanolophia canadaria, 164 
Hyperstrotia sp., 165 M. signataria, 164 
Hypna clytemnestra, 310 Mesoleuca ruficillata, 164 
Idaea demissaria, 164 Metanema determinata, 164 


Ideopsis guara, 56 M. inatomaria, 164 


318 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Metarranthis angularia, 164 P. troilus, 235 
Meteorus sp., 226 P. zelicaon, 203 
Michaelus vibidia, 70 Paradipsosphecia barbara, 119 
Microsynanthedon tanala, 120 Paramelia stygia, 225 
Mikania sp., 68 Pararge aegeria, 203 . 
Milionia aglaia, 275 Parasites of Lepidoptera, 40, 66, 69, 72, 
M. aroensis, 275 2268235 
M. basalis, 275 Pareronia valeria, 56 
M. callima, 275 Parthenocissus quinquefolia, 237 
M. diva, 275 Parthenos sylvia, 56 
M. dohertyi, 275 Passiflora costata, 141 
M. grandis, 275 P. mansii, 141 
M. isodoxa, 269 P. pittieri, 141 
M. mediofasciata, 275 P. serratifolia, 137 
M. paradesia, 275 Peromyscus leucopus, 196 
Milliera dolosalis, 120 Perrhybris lypera, 229, 230 
M. dolosana, 120 Pharneuptychia nr. pharnabazos, 200 
Miller, T. A., 207 Pheidole sp., 66 
Mitostemma glaziovii, 141 Phemiades milvius, 288 
Mutuuraia mysippusalia, 163 P. pohli, 288 
Narang, R. C., 112 P. pseudophineus, 286, 288 
Nathalis iole, 249 P. vergens, 288 
Nectandra gentlei, 230 Phocides pigmalion, 250 
Nematocampa limbata, 164 Phoebis agarithe, 248 
Nemora bistriaria, 164 P. sennae, 248 
N. rubrifrontaria, 164 P. statira, 149 
Neodiprion swainei, 40 Phyciodes batesii, 121 
Neptis hylas, 200 P. campestris, 121 
Nielsen, M. C., 157, 233 P. frisia, 247 
Nolina microcarpa, 304 P. phaon, 121 
N. texana, 304 P. tharos, 121 
Nymphalis antiopa, 265 Phycodes eucallynta, 119 
N. j-album, 265 Pieris brassicae, 133 
N. milberti, 265 P. callidice, 130 
Obituary, 62 P. protodice, 135, 174 
Ocotea sp., 229, 230 P. rapae, 133; (Artogeia), 266 
Oidaematophorus homodactylus, 163 Pinus kesiya, 275 
O. monodactylus, 163 Pithecoctenium crucigerum, 70 
Olethreutes albiciliana, 163 Plagodis alcoolaris, 164 
Oliver, C. G., 121, 153 P. fervidaris, 164 
Opler, P. A., 145 P. phlogosaria, 164 
Orphniospora atrata, 225 Plantago lanceolata, 40 
Orr, A. G., 54 Platt, A. P., 76 
Orthonama centrostrigaria, 164 Podisus maculiventris, 235 
Orthosia sp., 165 Polistes exclamans, 235 
Ostrinia nubilalis, 163 Polites coras, 266 
Oxylides faunas, 238 Polygonia interrogationis, 235, 265 
Palthis angulalis, 165 Polygonus leo, 250 
Panoquina panoquinoides, 249 Polyura spp., 56 
Papilio aristodemus ponceanus, 254 Predators of Lepidoptera, 35, 39, 40, 60, 
P. cresphontes, 248 148, 157, 183, 196, 235 
P. glaucus, 266 Probole ahenaria, Wee! 
P. helenus, 56 P. amicaria, 164 
P. iswara, 56 Propertius phineus, 279, 288 
P. nephelus, 56 P. propertius, 288 
P. polyxenes, 266 Prunus angustifolia, 22 


VOLUME 36, NUMBER 4 319 


P. maritima, 20, 28 S. melinus, 73, 247 

P. serotina, 20, 193 S. yojoa, 70 
Pseudolycaena dame, 71 Sylepta fluctuosalis, 163 
Pseudosphecia tenebrosa, 119 Synansphecia triannuliformis, 119 
Psychomorpha epimenis, 165 Synanthedon danieli, 120 
Pterocarpus sp., 71 Synchlora aerata, 164 
Pyrausta pertextalis, 163 Tarachidia erastrioides, 165 
Quercus falcata var. pagodaefolia, 227 Tebenna chrysostacta, 120 
Ragadia crisilda, 56 Telenomus prob. rileyi, 235 
Rahn, R., 158 Terinos terpander, 56 
Renia discoloralis, 165 Tetrastichus sp., 69 

R. factiosalis, 165 Thecla azaria, 73 


Rheedia edulis, 71 
Rhus copallina, 76 
Robbins, R. K., 65 


. crolus, 68 
. near enemia, 68 
. ericusa, 69 


Ps) ise lees | lh 


Samia cynthia, 198 . hemon, 71 
Sargent, T. D., 42 . hesperitis, 69 
Sassafras albidum, 193 . mathewi, 68 
Sbordoni, V., 185 Theobroma cacao, 71 
Scalarignathia kaszabi, 120 Thisbe irenea, 72 
Sceloporus graciosus, 151 Tmolus echion, 67 
Schweitzer, D. F., 18, 256, 303 Trogonoptera amphrysus, 56 
Scopula imboundata, 164 T. brookiana, 56 

S. inductata, 164 Tropidurus torquatus, 148 
Semiothisa bisignata, 164 Tryphera sp., 226 
Sesia bembeciformis, 120 Turnera panamensis, 71 

S. montelli, 120 Urbanus dorantes, 250 

S. okinawana, 120 } U. proteus, 250 
Shapiro, A. M., 174 Vaccinium caespitosum, 233 
Sicya macularia, 164. Vanessa atalanta, 265 
Siproeta stelenes, 246 V. cardui, 151 
Smith, D. S., 241 V. virginiensis, 265 
Solanum coconilla, 67 Vismia baccifera, 71 

S. lancaeifolium, 66 Waldbauer, G. P., 154, 192 

S. nudiflorum, 67 Waller, D. A., 178 

S. ochraceo-ferrugineum, 67 Williams, T. F., 76 

S. sanitwongsei, 67 Worth, C. B., 76 
Speyeria aphrodite, 266 Wylie, F. R., 269 

S. atlantis, 266 Xanthotaenia busiris, 56 

S. cybele, 266 Xanthorhoe ferrugata, 164 
Spilochalcis sp., 72 X. lacustrata, 164 
Stamp, N. E., 31, 290 Xanthotype urticaria, 164 
Sternberg, J. G., 154, 192 Xiphidium caeruleum, 70 
Stibochiona nicea, 56 Young, A. M., 155, 230, 310 
Stigmaphyllon lindenianum, 67, 69 Ypthima fasciata, 56 
Strymon basilides, 68, 69 Y. pandocus, 56 

S. bazochii, 158 Zale undularis, 165 

S. columella, 247 Zegris eupheme, 132 

S. martialis, 247 Zelkova drymeja, 145 


Zizyphus juazeiro, 202 


Lloyd M. Martin Memorial Research Fund, 320 


Journal of the Lepidopterists’ Society 
36(4), 1982, 320 


LLOYD M. MARTIN MEMORIAL RESEARCH FUND 


On 28 January 1982 a chapter in the annals of western Lepidopterology came to an 
end with the death of Lloyd Martin in Fresno, California. He was 69 years of age. 

In 1969 Lloyd retired from the Natural History Museum, where he had been curator 
of Lepidoptera for 33 years. He was a charter member of The Lepidopterists’ So- 
ciety and served our organization in many capacities, including a term as President 
in 1972. For more than 50 years he had been a member of the Lorquin Entomological 
Society, in which he held important offices for many years. 

Lloyd’s many friends and colleagues will miss his irreverent and earthy humor, 
his humanity, his devotion to the study of Southwestern Lepidoptera, and his con- 
stant support and encouragement of young entomologists. 

In tribute to his many significant contributions to the study of Lepidoptera, the 
Natural History Museum of Los Angeles County has established the “Lloyd M. 
Martin Memorial Lepidoptera Research Fund,’ to support the fields of inquiry to 
which he devoted most of his life and energies. Specifically, this fund will encourage 
field research in unexplored or little-known localities and the acquisition, preparation, 
and study of moths and butterflies of the western United States. 

Contributions to this Memorial Fund may be of any size, payable to “LACM Founda- 
tion” and mailed to my attention. 

JULIAN P. DONAHUE, Los Angeles County Museum of Natural History, 900 Exposi- 
tion Boulevard, Los Angeles, California 90007. 


Date of Issue (Vol. 36, No. 4): 18 March 1983 


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EDITORIAL STAFF OF THE JOURNAL 
THOMAS D. EICHLIN, Editor 


% Insect Taxonomy Laboratory 
1220 N Street 
Sacramento, California 95814 U.S.A. 


MAGDA R. PAppPp, Editorial Assistant 


DOUGLAS C. FERGUSON, Associate Editor THEODORE D. SARGENT, Associate Editor 


NOTICE TO CONTRIBUTORS 


Contributions to the Journal may deal with any aspect of the collection and study of 
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Literature Cited: References in the text of articles should be given as, Sheppard 
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the heading LITERATURE CITED, in the following format: 


SHEPPARD, P. M. 1959. Natural selection and heredity. 2nd. ed. Hutchinson, London. 
209 pp. 

196la. Some contributions to population genetics resulting from the study of 
the Lepidoptera. Adv. Genet. 10: 165-216. 


In the case of general notes, references should be given in the text as, Sheppard (1961, 
Adv. Genet. 10: 165-216) or (Sheppard 1961, Sym. R. Entomol. Soc. London 1: 23-30). 

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PRINTED BY THE ALLEN PRESS, INC., LAWRENCE, KANSAS 66044 U.S.A. 


CONTENTS 


HABITAT, DIVERSITY AND IMMIGRATION IN A TROPICAL ISLAND 
FAUNA: THE BUTTERFLIES OF LIGNUMVITAE KEY, FLORIDA. 


Dennis Leston, David Spencer Smith & Barbara Lenczew- 
Ski. UNS VA OS ON 


FIELD OBSERVATIONS OF FOODPLANT OVERLAP AMONG SYM- 
PATRIC CATOCALA Sone ON JUGLANDACEAE. Dale F. 
Schweitzer 


THE BUTTERFLIES OF KENT ISLAND, GRAND MANAN, NEW 
BRUNSWICK. G. David Maddox & Peter F. Cannell 


FLIGHT PATTERNS AND FEEDING BEHAVIOR OF ADULT MILI- 
ONIA ISODOXA PROUT AT BULOLO, PAPUA NEW GUINEA 
(GEOMETRIDAE). F. R. Wylie 


REDISCOVERY OF THE TYPE OF PAPILIO PHINEUS CRAMER AND 
ITs BEARING ON THE GENERA PHEMIADES HUBNER AND 
PROPERTIUS EVANS (HESPERIIDAE). Rienk de Jong 


SELECTION OF OVIPOSITION SITES BY THE BALTIMORE CHECK- 
ERSPOT, EUPHYDRYAS PHAETON (NYMPHALIDAE). Nancy E. 
SEQ oe 


GENERAL NOTES 


Field observations of divergent resting behavior among hickory feeding 
Catocala larvae (Noctuidae). Dale F. Schweitzer 


A late-season emergence of Callophrys (Sandia) macfarlandi (Lycaeni- 
dae). Richard Holland 


The correct placement of Everes ‘herrii.’ Richard A. Bailowitz 


Natural history of Hypna clytemnestra Cr. (Nymphalidae) in Costa Rica. 
Allen M. Young 


241 


256 


264 


269 


279 


290 


Volume 37 1983 Number 1 


ISSN 0024-0966 


JOURNAL 


of the 


LEPIDOPTERISTS’ SOCIETY 


Published quarterly by THE LEPIDOPTERISTS’ SOCIETY 


Publié par LA SOCIETE DES LEPIDOPTERISTES 
Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN 
Publicado por LA SOCIEDAD DE LOS LEPIDOPTERISTAS 


DEDICATED TO THE MEMORY OF 


Robert E. Silberglied 
1946-1982 


19 August 1983 


THE LEPIDOPTERISTS’ SOCIETY 


EXECUTIVE COUNCIL 


CHARLES V. COVELL, JR., President LINCOLN P. BROWER, 
CLIFFORD D. FERRIS, Vice President Immediate Past President 
ALBERTO Diaz FRANCES, Vice President JULIAN P. DONAHUE, Secretary 
CLAUDE LEMAIRE, Vice President RONALD LEUSCHNER, Treasurer 


Members at large: 


R. L. LANGSTON K. S. BROWN, JR. F. S. CHEW 
R. M. PYLE T. C. EMMEL G. J. HARJES 
A. M. SHAPIRO E. H. METZLER 


The object of the Lepidopterists’ Society, which was formed in May, 1947 and for- . 
mally constituted in December, 1950, is “‘to promote the science of lepidopterology in 
all its branches, .... to issue a periodical and other publications on Lepidoptera, to facil- 
itate the exchange of specimens and ideas by both the professional worker and the 
amateur in the field; to secure cooperation in all measures’ directed towards these aims. ; 

Membership in the Society is open to all persons interested in the study of Lepi- 
doptera. All members receive the Journal and the News of the Lepidopterists’ Society. : 
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members should send to the Treasurer full dues for the current year, together with their 
full name, address, and special lepidopterological interests. In alternate years a list of 
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numbers in each volume of the Journal, scheduled for February, May, August and 
November, and six numbers of the News each year. 


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4 
: 

Back issues of the Journal of the Lepidopterists’ Society, the Commemorative Vol- ' 
ume, and recent issues of the NEWS are available from the Treasurer. The Commem- 
orative Volume, is $6; for back issues, see the NEWS for prices or inquire to Treasurer. . 


Order: Mail to Ronald Leuschner, 1900 John St., Manhattan Beach, California 90266 } 
Lis 


: 
c. 
Journal of the Lepidopterists’ Society (ISSN 0024-0966) is published quarterly by the 
Lepidopterists’ Society, a non-profit, scientific organization. The known office of publi- 
cation is 1041 New Hampshire St., Lawrence, Kansas 66044. Second class postage paid | 
at Lawrence, Kansas, U.S.A. 66044. 
4 


Cover illustration: Adult of the squash vine borer, Melittia cucurbitae (Harris) (Sesi- 
idae), which occurs in the eastern half of the United States and along the Gulf Coast into 
Vera Cruz, Mexico. The larvae are destructive borers in the vines of various cultivars of 
Cucurbita spp. (squash, pumpkins and gourds). Original drawing by Dr. Charles S. Papp, 
Sierra Graphics & Typography, 1722 J Street #19, Sacramento, CA 95814, USA. 


JOURNAL OF 


Tue LeErPIpDOPTERISTS’ SOCIETY 


Volume 37 1983 Number 1 


Journal of the Lepidopterists’ Society 
37(1), 1983; 1-18 


TERRITORIAL BEHAVIOR OF NYMPHALIS ANTIOPA AND 
POLYGONIA COMMA (NYMPHALIDAE) 


ROYCE J. BITZER AND KENNETH C. SHAW 
Department of Zoology, Iowa State University, Ames, Iowa 50011 


ABSTRACT. The behavior of perching Nymphalis antiopa and Polygonia comma 
is described. Males of both species occupy and defend specific areas or territories. 
Territories of N. antiopa are much larger than those of P. comma. P. comma males, 
like Vanessa atalanta, patrol and defend the same areas. In contrast, N. antiopa males 
defend areas much larger than areas patrolled and change perch sites frequently. Ter- 
ritorial behavior of these two species and that of V. atalanta are compared and attempts 
are made to explain differences among the three species on the basis of varying degrees 
of competition for mating sites and females. 


Male butterflies adopt one of two general strategies for locating 
mates, perching or patrolling (Scott, 1974). Patrolling species search 
for females, whereas perching species perch on trees, bushes, herbs 
or the ground and wait for females to fly by. Many of the fast flying, 
relatively short-winged nymphalids are perching species (Joy, 1902; 
Shields, 1967; Baker, 1972; Dimock, 1978; Bitzer & Shaw, 1979(80)). 
Nymphalids and other perching butterflies vary in the degree of site 
attachment and in the intensity with which they pursue butterflies 
that pass near their perches. This has led to a controversy over wheth- 
er male butterflies show true territorial behavior (Baker, op. cit.; Scott, 
op. cit.; Silbergleid, 1977; Davies, 1978; Bitzer & Shaw, op. cit.). 

We have described territorial behavior in the nymphalid, Vanessa 
atalanta (L.) (Bitzer & Shaw, op. cit.). Territorial behavior in this 
species is characterized by 1) outlining territories by intermittent pa- 
trols, 2) perching on one or two specific sites within territories and 3) 
chasing all intruders, including conspecifics, butterflies of other species 
and birds. Conspecifics are chased vertically in a characteristic spiral, 
while other species of butterflies and birds are chased horizontally to 
the limits of the territorial boundary. In this study we describe ter- 
ritorial behavior of two other nymphalids, Nymphalis antiopa (L.) and 
Polygonia comma Harris. 


JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


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VOLUME 37, NUMBER 1 2) 


& 


Nymphalis antiopa 


TEWAM® (OVORRG I — SSS i eo re pe 


eee 


METERS 


ravine wall 


Fic. 1. Perching sites of N. antiopa males in ravine on 21 April 1979. Diagram 
shows southwest three-quarters of ravine: (@) perch sites at indicated times (CST); 
( ) total areas defended and patrolled by males identified by number; (+) site of 
interaction between adjacent males at indicated times; (-——) flight path of male #1 to 
site of interaction with male #3 and back to same resting spot; (----- ) boundary of 
territory #1 within boundary of territory #2. 


OBSERVATIONS 
Nymphalis antiopa 


N. antiopa males were observed in a ravine (46 m long, 28 m wide) 
in Railroad Park, Ames, Iowa (a wooded, hilly area, 51.8 hectares) on 
8 April 1977 and 14, 18 and 21 April 1979. Males entered the ravine 
between 1130 h and 1200 h (CST) and departed between 1530 h and 
1600 h. Males chased other N. antiopa and other species of butterflies 
and birds from territories averaging 308 + 124.7 m? (Table 1). Occu- 
pants flew short patrols every few minutes, outlining areas much 
smaller than the area defended (Table 1). After a patrol, males seldom 
returned to the same perch (Fig. 1). On 21 April, during 30 successive 
landings, a territorial male landed 0-11 m (x + S.D. = 3.06 + 2.36 m) 
from his previous perch, returning to the same perch only twice. 

On 18 and 21 April, males averaged 16.2 patrols/h (64 in 244 min) 
(Table 1) and 1.4% of their site occupation time (3.3 of 244 min) pa- 


4 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


NUMBER OF EVENTS 


Ye 
N. antiopa patrols P. comma interactions 
i N. antiopa interactions = V. atalanta interactions 


Fic. 2. Temporal distribution of activity of N. antiopa, P. comma and V. atalanta. 
N. antiopa—total number of patrols and interactions by single occupants for 14, 18, 
and 21 April 1979. P. comma—total number of interactions in a cluster of six perches 
and nine perches on 21 and 22 April 1979 respectively. V. atalanta—mean number of 
interactions/day for 10 days in mid-June 1979. 


trolling. Length of patrols ranged from 2-58 seconds (12.1 + 9.6 s) 
(Table 1). Frequency of patrolling was highest between 1315 h and 
1445 h (Fig. 2) and was positively correlated with frequency of in- 
truders entering the ravine. Eleven of 12 intreractions occurred be- 
tween 1310 h and 1450 h (Fig. 2). During the four days of observation, 
seven intruders entered the ravine from varying directions at an in- 
trusion frequency of 0.8/h (7 intrusions in 528 min). The four ravine 
occupants (one each day) chased the seven intruders 12 times for an 
interaction frequency of 1.80/h (Table 1). Mean interaction time was 
21.2 + 9.9 s (range 10-40 s) and occupants spent 0.6% (3.2 of 528 min) 
of their time interacting with intruders. 

Intruders entered the ravine flying about 5-6 m above the bottom 
of the ravine. The occupant flew from his perch, the angle depending 
upon the intruder’s location, to intercept the intruder. After meeting 
they spiralled together and climbed vertically 10-14 m (Fig. 3). After 
they leveled off, the occupant chased the intruder around one-half to 
three-quarters of a circle, 9-15 m in diameter (Fig. 3). When circling, 
the occupant was usually 30-50 cm behind the intruder. Three to four 
times during each chase the occupant closed to within a few cm of 
the intruder. The intruder eventually broke out of the circle and flew 
out of the ravine (Fig. 3). 

Because of the size of the ravine and the tendency for occupants to 
perch close to one end of the ravine, at times (3 times in 4 days) two 


VOLUME 37, NUMBER l| D 


a2 SS == 


Nymphalis antiopa 


5 10 
METERS 


Fic. 3. Flight pattern during typical interaction between two N. antiopa males: 
(——) flight path of territory occupant; (-—-—) flight path of intruder. 


males temporarily occupied the same ravine. On 18 April the original 
occupant at the southwest end of the ravine patrolled northeastward 
and flew through a second male’s territory. After six interactions (in 
12 min), during which the perching sites of the two males moved 
northeastward toward the end of the ravine, the original occupant 
drove the second male out of the ravine. On 21 April the original 
occupant drove out two different males that temporarily perched in 
the ravine (Fig. 1). One intruder perched on the side of the northeast 
end of the ravine at 1323 h. After one interaction at 1341 h he was 
driven high on the slope, where he set up a small territory. He was 
observed patrolling and occupying the indicated resting spot at 1402 
h. He remained in this new territory for at least 10 minutes, after 
which observations were concentrated on male #1. The same occu- 
pant drove another male out of the ravine after the intruder had perched 
for two minutes. In this interaction the occupant left his perch and 
flew about 18 m to the site of the perched intruder. When the intruder 
rose to meet the occupant, the intruder was driven from the ravine. 
The occupant apparently had spotted the intruder from his perch and 
eventually flew to investigate. Occupants also flew to investigate us 
or light-colored objects placed on a ledge near the bottom of the rav- 
ine. On 18 April the occupant flew out eight, 12 and 24 m from his 
perch to investigate us or the objects on the ledge. 

On 14 April 1979 a female entered the ravine from the southwest. 


6 JOURNAL OF THE.LEPIDOPTERISTS SOCIETY 


The male intercepted her, and the interaction was similar to a male— 
male interaction, until they had flown about one-third of a circle. The 
male prodded the smaller female, who then fluttered her wings twice, 
three seconds apart, each flutter lasting approximately one second. 
When she fluttered, the yellow bands on her wing margins were high- 
ly visible. After interacting for 20 seconds, the male prodded her 
downward, and they both dropped into long grass near the top of the 
ravine. Seven minutes later we found them copulating, and they stayed 
in copula approximately two hours. 

These observations suggest that N. antiopa males are unable to 
determine the sex of an intruder until an interaction is nearly over. 
By flying behind the intruder and prodding it, the male may be at- 
tempting to determine the intruder’s sex. If the intruder flips its wings 
the occupant recognizes it as a female and prods her to drop to the 
ground. The vertical climb allows the butterflies to rise above the tree 
canopy to circle. 

Besides chasing conspecifics, perching N. antiopa males also chased 
birds and falling leaves. N. antiopa is more discriminating than V. 
atalanta (Bitzer & Shaw, op. cit.) when chasing falling leaves. Al- 
though the wind blew many leaves into the ravine, the butterflies 
chased only those which drifted with a rocking motion at approxi- 
mately the velocity of an intruder. On 8 April 1977 a perching male 
followed about 30 cm behind one of these leaves and prodded it two 
or three times before the latter landed on the ground. It is advanta- 
geous for a butterfly which perches in early spring not to waste time 
and energy chasing the large number of fallen leaves which blow 
about at this time. The butterflies did not discriminate between birds, 
however, and chased 98% of those which flew over. Their pursuits of 
birds suggest that N. antiopa males can adjust their angle of climb so 
as to intercept objects passing overhead at different velocities. Birds 
were chased horizontally until they left the ravine. 


Polygonia comma 


P. comma males of the Spring brood were observed on 27 March 
1978 and 21 and 22 April 1979 in Railroad Park, Ames, Iowa. Males 
perched on fixed bare spots on the ground or on the sunlit sides of 
trees about 1-2 m above the ground. They began perching between 
1515 h and 1530 h (CST) and left between 1715 h and 1800 h (Fig. 2). 
Earlier in the day (1200-1500 h) butterflies were seen flying up and 
down the sides of several ravines in the area. 

Like V. atalanta (Bitzer & Shaw, op. cit.), P. comma males occupied 
one or a few fixed resting spots (Fig. 4). Unlike V. atalanta, they did 
not consistently outline a fixed territory by patrolling. On 21 April 


VOLUME 37, NUMBER 1 ie 


4 


0) 5 10 
METERS 


Polygonia comma 


~ ae 
+ 


/ 
if ata \ ® 


p / 


/ 
\ 
\ 
/ 


+ \ 3 
\ 2 e 
° ® 
7 + 
7 
> 7 
7 
~ 7 
BSE 7 @B \ 
ve oss + ® 


ravine 


v 


Fic. 4. Territories and perch sites of P. comma males on wooded hilltop, 21 and 
22 April 1979: ( ) areas patrolled by occupants or areas from which intruders chased; 
(@) perch sites; (®) trunk sites of canopy trees; (+) shrubs or 1-2 m saplings; (-—-) 
flight paths of intruders; (Ravine) top of south-east side of ravine in Fig. 1. 


only one of the six butterflies patrolled the area around its resting 
spots (P2, 3 times), while on 22 April four of 10 butterflies patrolled 
fixed areas (Fig. 4). The areas outlined for the remaining butterflies 
in Fig. 4 indicate approximate areas from which passing intruders 
were chased. Butterflies rested on the same spots on 21 and 22 April. 
After one butterfly displaced another from his perch, the new occu- 


8 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


pant used the same perch. This suggests that P. comma males, like 
V. atalanta (Bitzer & Shaw, op. cit.) choose perches by orienting to 
certain visual features of the environment, including light-dark con- 
trast. 

Intruders always entered from the north-northwest, and the simi- 
larity of flight paths suggests that these butterflies navigate along im- 
portant visual features. On 22 April 18 of 24 patrolling butterflies flew 
along part or all of one of the three paths shown in Fig. 4, and four 
intruders went through all territories from P6 through P7 to T4. Al- 
though no data were recorded for 21 April (because so many butter- 
flies were interacting simultaneously), most of the intruders passed 
through P6 and P2 while occupants chased them. Approximately 90% 
of these intruders continued through P1 and P7, while the remainder 
flew through P1 and P4 (Fig. 4). 

Most interactions occurred between 1615-1645 h (Fig. 2), and the 
number of interactions varied among perches. Of six perches occu- 
pied on 21 April, eight interactions occurred in P1, 13 in P2, one in 
P3, none in P4, three in P6 and one in P7. These 26 interactions were 
observed in 87 minutes for a mean frequency of 17.9/h for the cluster 
of six perches. The greatest frequency for the cluster in a 15 minute 
period was 36/h between 1615 h and 1630 h. Of nine perches occu- 
pied on 22 April, 14 interactions occurred in P2, none in P3, one in 
P4, one in P6, none in P7, six in Tl, one in T2, one in T4, and none 
in T6. These 24 interactions were observed in 154 minutes, giving a 
mean frequency of 9.4/h for the cluster. The greatest mean frequency 
in a 15 minute period was 16/h between 1645 h and 1700 h. The mean 
interaction frequency per hour per territory was 4.9 + 4.4 (Table 1). 
Time of occupancy for sites with the highest frequency of interactions 
(Pl, P2, Tl) ranged from 35-115 minutes. Range of occupation 
for other perches was 0-30 minutes. Perches with the least number 
of interactions (P3 and P4) were not along the main intrusion routes. 

We observed two types of P. comma interactions (Fig. 5). In 49 of 
50 interactions the occupant dashed up at the intruder, flying about 
1-2 m above the ground, and both butterflies spiraled tightly together, 
rising at a 50-60° angle to a height of 6-8 m in 3-5 seconds (Fig. 5A). 
Then the occupant began to chase the intruder up a half-circle, 40- 
50 m in diameter, rising at a shallower angle to 15-30 m. After one 
butterfly broke out of the circle, the other descended to the resting 
spot in a twisting path. We timed eight of these interactions, and they 
ranged from 12—40 seconds (x = 24 + 8.8 s). | 

In the other type of interaction the occupant intercepted the in- 
truder and both spiralled to a height of about 9 m before levelling off 
(Fig. 5B). Then one butterfly, 30 cm ahead of the other, dropped 


VOLUME 37, NUMBER 1 9 


METERS 


Polygonia comma 


) flight 


Fic. 5. Flight patterns during interactions of pairs of P. comma males: ( 
path of territorial occupant; (-——) flight path of intruder. A, typical interaction; B, 
single atypical interaction. 


abruptly to about 2 m. Again the two spiralled and rose, now to about 
6 m, then dropped again, this time just missing the trunk of a large 
tree. Once past the tree one butterfly chased the other along a straight 
line until both flew out of sight. Five seconds later and 30 seconds 
after the intruder had arrived, a butterfly flew back from the direction 
they had gone and landed on the perch. Baker (op. cit.) observed 
similar interactions in the nymphalids Inachis io (L.) and Aglais ur- 
ticae (L.). 


DISCUSSION 


Based on Brown’s (1975) definition of a territory as a “fixed area 
from which intruders are excluded by some combination of adver- 
tisement ..., threat and attack,” males of N. antiopa and P. comma 
occupy territories and exhibit territorial behavior. N. antiopa differs 
from P. comma and V. atalanta (Bitzer & Shaw, op. cit.) in that the 
area patrolled is not equivalent to the area defended. The behavior 
of the above three species is compared in Table 1, which is the basis 
for the following discussion. The differences among the three species 
may reflect varying degrees of competition for mating sites and fe- 
males among the three species. 

According to Huxley’s (1934) “elastic disc theory,’ increasing com- 
petition for territories should decrease the area which can be suc- 
cesstully defended. N. antiopa males roamed over relatively large 


10 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


areas of the occupied ravine, suggesting that territories were not in 
short supply. This was supported by the relatively low frequency of 
intruders and interaction frequencies, as well as the relative ease at 
which intruders were dislodged. This is in contrast to V. atalanta 
males which had much higher interaction frequencies and occasion- 
ally split territories with intruding males (Bitzer & Shaw, op. cit). 

Although they had the lowest interaction frequency, N. antiopa 
shows the highest mean frequency of patrolling. The latter may have 
been effected by the larger area which they defended and the variety 
of directions from which females and intruders entered the territories. 
The relatively large size of N. antiopa territories should increase the 
possibility that a female will fly through (Baker, op. cit.). Continually 
changing perches also may be correlated with the need to defend such 
larger territories and to spot potential mates. The relatively small areas 
patrolled and the shortest mean duration of patrols may be means of 
countering the energy demands of frequent patrols. 

P. comma and V. atalanta males probably exhibited lower territory : 
male ratios (Baker, op. cit.) than N. antiopa. There were a greater 
number of P. comma males/unit area, and the territories (=areas pa- 
trolled and/or defended) were smaller. Although territories of V. ata- 
lanta are intermediate in size, there were highly desired or optimal 
territories that were always occupied night after night and year after 
year and were reoccupied within five seconds to five minutes after 
experimental removal of occupants (Bitzer & Shaw, op. cit.). The large 
number of P. comma males per unit area supports competition for 
territorial sites, but it is difficult to understand why some apparently 
premium sites (e.g., Tl and Pl, Fig. 4) were not occupied both days 
of observation. The small size of P. comma territories apparently ne- 
gated the need for patrols. There were patrols in only five territories 
in two days of observation and 17 of 38 patrols occurred in territory 
T1, the largest territory (Fig. 4). 

In addition to greater competition increased compactness of terri- 
tories and fixity of perch sites also could be effected by increasing 
tendency for intruders to enter defended areas along specific flight 
paths. Constancy of flight paths could be the result of at least two 
factors, prominent visual features and tendency for females to fly 
through as many male territories as possible. Baker (op. cit.) suggests 
that the nymphalids Inachis io and Aglais urticae establish mating 
territories along visual lines of demarcation, such as hedges, walls 
and rows of trees. Optimal territories (those most frequently occupied 
and with the highest frequency of interactions) of V. atalanta oc- 
curred where sidewalks intersect and near sides of buildings on the 
Iowa State University campus (Bitzer & Shaw, op. cit.) and along the 


VOLUME 37, NUMBER 1 I 


tree-lines of forest margins in more natural habitats (Bitzer & Shaw, 
unpublished observations). Females of polygynous species have 
greater investment in offspring than males and should be very selec- 
tive of mates (Trivers, 1972). The opportunity to fly through a number 
of male territories, possibly two or more times, along a given flight 
path (Fig. 4) should greatly enhance a female’s ability to choose a 
genetically superior male, possibly expressed by his ability to occupy 
a highly contested territory, his chasing vigor or endurance, and/or 
the nature of his aphrodisiac (see discussion of the possibility that 
males of polygynous insect species have individual “signatures ’ which 
females can use to identify them during a testing period; Lloyd, 1981). 
If females do fly along specific, visually demarcated lines and through 
as many territories as possible, there should be intense competition 
for territories along these lines. Non-territorial males would be ex- 
pected to fly the same paths, assessing, during their flight interactions 
with territorial males, the feasibility of attempting to displace a ter- 
ritory holder or of establishing a new territory. 

When competition is intense males would benefit if they could rec- 
ognize females early in an interaction. The observation of one male-— 
female interaction suggests that N. antiopa males may not recognize 
an intruder’s sex until late in the interaction (approximately 20 s after 
the beginning of the interaction). In contrast, V. atalanta males may 
be able to discern an intruder’s sex within five seconds after first 
encountering an intruder. When interaction frequencies are high and 
there is an increased risk that another intruder will occupy his terri- 
tory before he can drive the current intruder way, a V. atalanta male 
will cut the interaction short after a few seconds of hovering by driv- 
ing him horizontally to the edge of his territory (Bitzer & Shaw, op. 
cit.). One suspected male-female interaction supports rapid sex rec- 
ognition. After a few seconds of hovering with an intruder, the two 
butterflies dropped downward through a bush (a move not unlike that 
of the N. antiopa male and female; Bitzer and Shaw, unpublished 
observations). Unfortunately, an extensive search failed to uncover 
the pair. 

A number of perching species, including some nymphalids, contin- 
ually change perches, and this has led some investigators to doubt 
whether any perching butterflies are territorial (Scott, op. cit.). If males 
change perches this suggests that there is no selective advantage to 
occupying a fixed perch. N. antiopa males defend areas larger than 
they patrol; therefore, it is advantageous for them to change perches 
within the defended area. Since mating territories of many nympha- 
lids are chosen, not for female resources they contain, but because 
they possess prominent visual markers affecting female flight paths 


Wy) JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


(Baker, op. cit.; Bitzer & Shaw, op. cit.), lack of prominent visual 
markers would negate males occupying fixed perches. Shields (op. 
cit.), in his study of hilltopping species, reported that three species 
of Vanessa, including V. atalanta, changed perches frequently. A 
published photograph of the study area indicates scrub vegetation 
showing little variation in height and apparent lack of easily differ- 
entiated visual lines of demarcation. Scott (op. cit.) reports four species 
of butterflies with different mate-locating behaviors in different parts 
of their ranges. Males from different populations of two species chose 
different topographic features for perching sites; in the other two 
species, some populations perched while others only patrolled. 
Whether a species occupies, defends and/or patrols fixed areas may 
be conditional upon local ecological conditions, such as availability 
of easily demarcated visual lines, predation pressure, and density of 
conspecific and interspecific competitors. 

If ecological conditions affect fixity of perch sites between species 
and between populations of the same species, could they affect flight 
patterns used by occupants to pursue intruders? In our earlier paper 
(Bitzer & Shaw, op. cit.) we suggested that the vertical spiral helix of 
V. atalanta facilitated the occupant dropping quickly back into his 
territory to pursue other intruders, while possibly disorienting the 
intruder in the overhead canopy. In contrast the series of dives and 
climbs of I. io and A. urticae (Baker, op. cit.) seem more adapted to 
driving the intruder up to 200 m from his territory in open country. 
Baker showed that the distance that intruders are driven from the 
territory is a compromise between the distance necessary to reduce 
the chance of the intruder returning and the time required for the 
owner to return before his territory is occupied by another conspe- 
cific. The flight pattern of N. antiopa (Fig. 3) appears adapted for 
driving the intruder to the edge of the ravine and out of sight of the 
ravine bottom. The two flight patterns of P. comma suggest that this 
species may be able to adapt its flight pattern based upon the nature 
of the tree canopy and the intensity of competition for perching sites. 
Forty-nine of 50 occupant-intruder interactions involved driving the 
intruder above the canopy. However, one interaction was very similar 
to that of I. io and A. urticae, which characterize butterflies of more 
open terrain. 

Interspecific competition for perching sites, such as occurs in hill- 
topping species (Shields, op. cit.), may result in selection for species 
to mate at different times of the day. Otherwise, perching butterflies 
would expend considerable energy chasing intruders of other species. 
Pairs of the three species reported on here may have undergone such 
selection. The territorial periods of N. antiopa and P. comma and 


VOLUME 37, NUMBER 1 1} 


those of P. comma and V. atalanta overlap slightly, i.e., when terri- 
torial activities are just beginning or just terminating (Fig. 2). N. an- 
tiopa’s preference for ravines also may spatially isolate them from P. 
comma. However, one observation emphasizes the importance of 
temporal isolation between these two species. N. antiopa and P. com- 
ma males occupied the same area on different days, a level area near 
the top of a ravine. 


LITERATURE CITED 


BAKER, R. R. 1972. Territorial behavior of the nymphalid butterflies Aglais urticae 
and Inachis io. J. Anim. Ecol. 4:453-469. 

BITZER, R. J. & K. C. SHAW. 1979(80). Territorial behavior of the Red Admiral, Va- 
nessa atalanta (L.) (Lepidoptera: Nymphalidae). J. Res. Lepid. 18:36—49. 

BROwN, J. L. 1975. The evolution of behavior. W. W. Norton and Company, Inc., 
New York. 

Davies, N. B. 1978. Territorial defense in the speckled wood butterfly (Pararge 
argia): The resident always wins. Anim. Behav. 26:138-147. 

Dimock, T. E. 1978. Notes on the life cycle and natural history of Vanessa annabella 
(Nymphalidae). J. Lepid. Soc. 32:88-96. 

HUXLEY, J. S. 1934. A natural experiment on the territorial instinct. British Birds 27: 
270-277. 

Joy, N. H. 1902. No title. (Fighting of males of Apatura iris L.) Proc. Entomol. Soc. 
Lond. 19:x]-xli. 

LLoyp, J. A. 1981. Sexual selection: individuality, identification, and recognition in 
a bumblebee and other insects. Florida Entomol. 64:89-118. 

ScoTT, J. A. 1974. Male locating behavior in butterflies. Am. Midl. Nat. 91:103-117. 

SHIELDS, O. 1967. Hilltopping. J. Res. Lepid. 6:69-178. 

SILBERGLIED, R. FE. 1977. Communication in the Lepidoptera. Pages 362-402, in T. 
A. Sebeock (ed.). How animals communicate. Indiana Univ. Press, Bloomington. 

TRIVERS, R. L. 1972. Parental investment and sexual selection. Pages 136-179, in B. 
Campbell (ed.). Sexual selection and the descent of man. Aldine, Chicago. 


Journal of the Lepidopterists’ Society 
37(1), 1983, 14-17 


A NEW SPECIES OF AGROTIS OCHS. (NOCTUIDAE) 
FROM SABLE ISLAND, NOVA SCOTIA 


KENNETH NEIL! 


Department of Biology, Dalhousie University, 
Halifax, Nova Scotia, Canada B3H 4J1 


ABSTRACT. A new species of Agrotis Ochs. from Sable Island, Nova Scotia is 
figured and described. 


Recent biological surveys of Sable Island, Nova Scotia by the staff 
of the Nova Scotia Museum have added much to the local knowledge 
of the Lepidoptera of the island. One of the more interesting captures 
taken during these studies was a small series of pale Agrotis Ochs. 
resembling Agrotis volubilis Harv. and Agrotis stigmosa Morr. (for- 
merly Agrotis volubilis f£. stigmosa Morr.). Subsequent investigation 
of these specimens plus additional material collected by the staff of 
the Biosystematics Research Institute, Ottawa, during the summer of 
1967 showed that they represented an undescribed species with dis- 
tinguishing characters in the female genitalia. 

Agrotis volubilis occurs from Newfoundland (Morris, 1980) and Nova 
Scotia (Ferguson, 1954) south to North Carolina, west to the Pacific 
(Forbes, 1954). Agrotis stigmosa occurs from Massachusetts west to 
North Dakota (Ahmandi, 1979). The early stages and host plants of 
A. stigmosa are unknown. The larva of A. volubilis has been reared 
from Achillea millefolium L. (McCabe, 1981). 


Agrotis arenarius, new species 


Description. Upperside of forewing overall much lighter and with lines and mark- 
ings less distinct than in volubilis and stigmosa. Ground of forewing light sandy brown. 
Costal area slightly darker with some light-grey scaling along R and Cu. A dark 
“W”’-shaped patch present in terminal area opposite cell. Subterminal area with a series 
of dark, elongate, “V-shaped marks between the veins. Reniform and orbicular spots 
concolorous with costa, overlain with light-grey scales and brown annuli. Reniform 
outlined on inner and outer edges with a narrow band of black scales. Orbicular with 
a similar outline on posterior half. Area between reniform and orbicular and on outer 
edge of reniform darker. Basal dash and claviform spot fused, outlined with black and 
filled with dark brown. Postmedial and antemedial lines present but faint and indis- 
tinct, most readily visible as light-brown patches at costa. Fringe concolorous. 

Upperside of hind wing lighter in male, mainly dirty white with a fuscous border. 
Hind wing of female more suffused with fuscous overall. Veins and discal spot in both 
sexes deanened with darker brown scales. Fringe white. 

Underside of forewing with markings as in stigmosa and volubilis but much ies 
lacking the brownish shades present in those species. Ground of undersides light sandy 
brown. A darker shade present in cell extending from base to postmedial line. Subter- 
minal and postmedial lines darker and distinct. Discal spot blackish. 


1 Current address: Department of Biological Sciences, Simon Fraser University, Burnaby, British Columbia, Can- 
ada V5A 1S6. 


VOLUME 37, NUMBER 1 15 


Fics. 1-6. Agrotis spp.: 1, A. arenarius 2, holotype; 2, A. arenarius ¢, allotype; 
3, A. stigmosa 2, Chatham Lab., Light Trap, 28 May 1935; 4, A. stigmosa ¢, Brooklyn, 
Long Island, New York, 13 May 1903; 5, A. volubilis 2, Kentville, King’s Co., Nova 
Scotia, 24 May 1979; 6, A. volubilis 3, Sheffield Farm near Canning, King’s Co., Nova 
Scotia, 10 June 1980. All about x2. 


Underside of hind wing dirty white. Costal area darker. Discal spot blackish. 

Vestiture of head and body as in stigmosa and volubilis. No visible differences in 
antennae, palpi, or other external structures. 

Length of forewing: males, 13.8-17.7 mm; females, 16.7-18.5 mm; holotype female, 
18.3 mm; allotype male, 17.2 mm. Mean wing length: male paratypes (10), 17.44 mm; 
female paratypes (6), 17.83 mm. 

Male genitalia (Figs. 7-9). Identical to those of Stigmosa and volubilis. In some 
specimens of arenarius the valve is more convex apically, but this is a variable character 
and cannot be used to separate arenarius from stigmosa and/or volubilis. 

Female genitalia (Fig. 10). Appendix bursae shorter in arenarius than in both 
stigmosa (Fig. 12) and volubilis (Fig. 11). Female genitalia otherwise similar. 

Types. HOLOTYPE: 2, Henry House, Sable Island, Nova Scotia, 2 July 1980, E. 
Quinter (Fig. 1). ALLOTYPE: ¢, Same data as holotype but taken 4 July (Fig. 2). PARA- 
TYPES: 1 3d, 1 2, same data as holotype but taken 3 July 1980; 1 ¢, 1 2, same data as 


16 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Fics. 7-12. Genitalia of Agrotis spp.: 7, A. arenarius 6, allotype, aedeagus omit 
ted; 8, A. arenarius 6, right valve of paratype, Sable Island, Nova Scotia; 9, A. are- 
narius 36, aedeagus of allotype; 10, A. arenarius °°, holotype; 11, A. volubilis 2, 
Sheffield Farm, King’s Co., Nova Scotia; 12, A. stigmosa 2, Chatham Lab., Light Trap. 


VOLUME 37, NUMBER l We 


holotype; 2 66, same data as allotype; 4 66, 2 2°, same data as holotype, 3 66, 1 
2, Meteorological Station, Sable Island, Nova Scotia, 15 July 1977, B. Wright; 1 @, 
West Light, Sable Island, Nova Scotia, 25 July 1976, B. Wright. Holotype and allotype 
have been deposited in the American Museum of Natural History collection. Paratypes 
have been deposited in the Canadian National Collection, Nova Scotia Museum, and 
the K. Neil collection. 

Distribution. This species is known only from Sable Island, Nova Scotia. 

Early stages. Unknown, but appears to be associated with Ammophila breviligulata 
Fern., as adults have been taken most commonly where the grass is most abundant on 


Sable Island. 


REMARKS 


Arenarius represents the second endemic moth to be recorded from 
Sable Island, the first being Orgyia leucostigma sablensis Neil and 
like that species, represents a Pleistocene relict which survived gla- 
ciation on the offshore refugium of which Sable Island is the last trace. 
Adults have been taken from early June until late July. 


ACKNOWLEDGMENTS 


I thank Barry Wright of the Nova Scotia Museum for his many helpful suggestions 
and comments made during the preparation of this paper, for providing material from 
the Nova Scotia Museum collection, and for reviewing the final manuscript; Eric Quin- 
ter for providing additional specimens of arenarius; Dr. J. D. Lafontaine of the Bio- 
systematics Research Institute, Agriculture Canada, Ottawa, for providing specimens 
of arenarius and stigmosa from the Canadian National Collection; and Mary Primrose 
of Dalhousie University for photographing the types. 


LITERATURE CITED 


AHMADI, A. A. 1977. A revision and review of the North American species of Agrotis 
and Feltia known to occur north of the American border (Lepidoptera, Noctuidae). 
Ph.D. thesis, Comell University, Ithaca, New York. 

FERGUSON, D. C. 1954. The Lepidoptera of Nova Scotia: (Macrolepidoptera). N.S. 
Mus. Sci. Bull. 3:214 pp. 

FORBES, W. T. M. 1954. The Lepidoptera of New York and neighboring states. Pt. 
III. Cornell Univ. Agr. Expt. Sta. Mem. 329:433 pp. 

McCaBgE, T. L. 1981. The larva of Agrotis volubilis Harvey (Lepidoptera: Noctuidae). 
J. N.Y. Entomol. Soc., 89(2):59-64. 

Morris, R. F. 1980. Butterflies and moths of Newfoundland and Labrador; the Mac- 
rolepidoptera. Agric. Can. Publ. 1691:407 pp. 


Journal of the Lepidopterists’ Society 
37(1), 1983, 18-23 


A NEW SPECIES OF SCHINIA (NOCTUIDAE) 
FROM MANITOBA AND SASKATCHEWAN WITH 
DESCRIPTION OF ITS LIFE HISTORY 


D. F. HARDWICK 
Biosystematics Research Institute, Ottawa, Ontario, Canada KIA 0C6 


ABSTRACT. Schinia verna, closely related to S. honesta (Grote), is described as 
new. The new species is a resident of the parkland belt of Manitoba and Saskatchewan 
and feeds in the larval stage in the flowering heads of Antennaria neodioica Greene. 
The immature stages of the new species are described. 


While undertaking field work in the Glenboro area of southern 
Manitoba in the spring of 1979, my wife and I collected and reared a 
new species of Schinia, closely related to Schinia honesta (Grote, 
1881). The following year a specimen of the undescribed species, 
collected at Saskatoon, Saskatchewan, was submitted to the Biosys- 
tematics Research Institute, Agriculture Canada, Ottawa, for identi- 
fication. 


Schinia verna, new species 


(Eiicisamlee2) 


Description. Eyes greatly reduced, elliptoid. Antennae filiform in both sexes. Fore- 
tibia with a conspicuous, elongate apical spine on inner side and a shorter apical spine 
on outer side; two or three additional slender, inconspicuous spines on either side of 
foretibia proximal to apical spines. 

New species smaller and paler than Schinia honesta (Fig. 3), which it most closely 
resembles in maculation. New species with dark areas of wing suffused with reddish 
and with pale areas more extensive than in honesta. 

Vestiture of head and thorax hair-like, light grey suffused with mauve; abdomen 
black-scaled with an overlay of pale-grey hair. Underside of body white. 

Forewing olive-brown, heavily suffused with mauve or reddish-brown and marked 
with light grey and white. Reddish suffusion lost in worn specimens. Transverse an- 
terior line indicated only by color change, roughly triarcuate, the median arc very broad 
and deep; often an elongate narrow notch anterior to median arc. Basal space olive- 
brown, suffused with reddish; pale grey at immediate base. Transverse posterior line 
sinuous, evident as a grey shade or indicated only by color change. Median space white, 
shaded with grey posteriorly and often with a narrow grey band along costal margin. 
Orbicular spot prominent, circular. Reniform spot only slightly larger than orbicular; 
both spots concolorous with basal space. Subterminal line white, forming an inward 
notch opposite cell and another toward anal angle. Subterminal space concolorous with 
basal space, often shading to grey proximally. Terminal space pale grey. Fringe grey 
with white points. 

Hind wing black, with a white central area containing a large black discal spot. Some 
pale shading in black outer-marginal area. Fringe white with a darker basal shade. 

Underside of forewing white with a black basal dash, black reniform and orbicular 
spots and black and grey subterminal band. Fringe white with dark intervenal dashes. 
Underside of hind wing white with a dark-grey discal spot and dark-grey patch near 
anal angle; suffused with grey basally. Fringe white. 

Expanse: 19.9 + 1.6 mm (16 specimens). 


VOLUME 37, NUMBER Il 19 


Fics. 1-6. Schinia spp.: 1 & 2, S. verna, n. sp., holotype and allotype, Glenboro, 
Manitoba; 3, S. honesta (Grote), Monument Pk., Linn Co., Ore.; 4, S. verna ovipositing 
in head of Antennaria neodioica Greene; 5, S. verna, female genitalia; 6, S. verna, 
male genitalia. 


Male genitalia (Fig. 6). Indistinguishable from those of honesta (see Hardwick, 1958, 
p. 69), except for their proportionately smaller size. 

Female genitalia (Fig. 5). Very similar to those of honesta and differing chiefly in 
the conformation of the ovipositor valve, which is broadly rounded apically in verna 
rather than pointed as in honesta. 


20 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Type material. HOLOTYPE: 6, Glenboro, Manitoba, 11 June 1979, D. & V. Hardwick. 
ALLOTYPE: @, same locality and collectors, 6 June 1979. PARATYPES: 8 66, 6 22, 
Glenboro, 9 to 12 June 1979, D. & V. Hardwick; 1 6, Saskatoon, Saskatchewan, 23 
May 1980. The type material is in the Canadian National Collection (Type No. 16876). 


Life History and Habits 


The population of Schinia verna trom which the type series was 
taken is located in south-central Manitoba, north of Glenboro, in an 
area which has recently been incorporated into Spruce Woods Pro- 
vincial Park. The meadow which constitutes the habitat of the species 
here is in a semi-wooded region of spruce and aspen. It was formerly 
part of the farm of Mr. Wm. Shewfelt and is still grazed by his cattle; 
the meadow supports a variety of spring-blooming plants. 

Two species of Antennaria (pussy toes), are present in the area, A. 
neodioica Greene and A. aprica Greene. Eutricopis nexilis Morrison, 
a heliothentine that feeds on a number of species of Antennaria from 
eastern to western North America, was flying abundantly among the 
patches of pussy toes. The first few specimens of S. verna were netted 
and killed on the assumption that they were also E. nexilis. Although 
eggs and larvae of S. verna were only found in the heads of Anten- 
naria neodioica, partially grown larvae would also readily accept the 
heads of A. aprica as food. 

The very large egg of verna is deposited (Fig. 4) deeply within the 
flowering head of the food plant. The newly hatched larva usually 
burrows directly to the achene layer and feeds on the seeds and also 
on the tissue of the receptacle. The early instars are very cannibalistic, 
and a number of dissected heads contained the remnants of both S. 
verna and E. nexilis larvae as well as a healthy first or second stadium 
verna larva. First instars hatching in previously occupied heads have 
the habit of crawling out of the head and seeking another to enter. 
The survival rate among these small wanderers is probably not very 
high. In most cases observed the displaced first instars could not pen- 
etrate a second head and fell from the food plant. 

Second instars seem quite capable of migrating from one head to 
another. Commonly, larvae remain in the initial head until reaching 
the second stadium, at which time they seek another, entering the 
second head from the top. Second stadium and early third stadium 
larvae continue to feed within the head, and their habit of attacking 
the receptacle as well as the seeds usually causes a complete disin- 
tegration of the head with the florets and pappus falling free. 

Late third instars and subsequent stadia feed from outside the head. 
Third and fourth instars often tie adjacent Antennaria heads together 
to form a protective shelter from which they feed; on becoming pre- 


VOLUME 37, NUMBER 1 2) 


Fics. 7-10. Schinia verna, n. sp., on Antennaria neodioica Greene; 7, molting nest 
of fourth stadium larva; 8, same, opened to show larva; 9, ultimate stadium larva, 
lateral; 10, same, dorsal. 


molt, they form a very definite nest of floral parts (Figs. 7, 8) in which 
they can remain quiescent until moulting is completed. 

Last instars feed exposed, attacking the base of the head from a 
position on the stem (Fig. 10). Larvae of Schinia verna mature in five 
stadia and require a mean period of 17.1 days from hatching to the 


D9; JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


cessation of feeding to complete their development. The mature larva 
retires to the ground and digs a short tube below the surface at the 
end of which it forms its pupal cell. The species is univoltine, the 
pupae remaining in diapause until the following spring. 


Immature Stages 


Egg. Very large. Translucent white when deposited. Showing little change until two 
days after deposition, when anterior end becomes suffused with pink. On day of hatch- 
ing pink suffusion fades, and mandibles become visible at micropylar end. All eggs 
observed had an incubation period of three days. 

First instar. Head, prothoracic and suranal shields black. Trunk dirty-white with 
small black setal bases. Mean duration of stadium, 3.1 days. 

Second instar. Head black. Prothoracic and suranal shields somewhat paler. Trunk 
medium greenish-grey, becoming paler toward end of stadium. Setal bases black, pro- 
portionately much larger than in first stadium. Mean duration of stadium, 2.6 days. 

Third instar. Head capsule black. Prothoracic and suranal shields dark brown. Trunk 
medium grey with large black setal bases. Mean duration of stadium, 2.5 days. 

Fourth instar. Head black with a large brown patch on either side; often a whitish 
triangle on frons. Prothoracic shield black with a yellowish transverse median band. 
Suranal shield black with an anchor-shaped yellowish median mark. Trunk medium 
grey with a variably expressed, yellow, transverse median shade on each segment, 
yellow shading becoming more intense toward end of stadium. Setal bases large, black. 
Rims of spiracles dark brown. Mean duration of stadium, 3.3 days. 

Fifth instar (Figs. 9, 10). Head fawn dorsally and laterally and finely mottled with 
light brown; frons dull white; a black inverted V through adfrontal areas; a pair of black 
spots above inverted V and a black patch in ocellar area. Prothoracic shield black with 
a pale-yellow transverse median band. Suranal shield undistinguished from remainder 
of trunk. Trunk uniform greyish-white. A variably expressed transverse yellow shade 
on dorsum of each segment. Setal bases very large and black, giving larva a checkered 
appearance. Spiracles small, dark brown. Mean duration of stadium, 5.6 days. 

Pupa. Well sclerotized; orange-brown, without green suffusion across appendages. 
Mesothoracic legs relatively long, terminating only a short distance anterior to apex of 
proboscis. Metathoracic legs evident as triangulate plates distal to apex of proboscis. 
Anterior third of each of abdominal segments 5 to 7 not raised above remainder of 
segment; anterior third of these segments finely and only sparsely pitted and not more 
darkly pigmented than remainder of segments. Spiracles set into shallow oval pits; rims 
of spiracles elevated to form short but definite tubes. Cremaster reduced to 2, rather 
stout, usually curved bristles, borne at apex of a conical prolongation of 10th abdominal 
segment. 


Remarks 


The spring of 1979 was a very late one in central Canada, and the 
flight data for the type series may indicate a period of activity that is 
later than normal. The type locality was revisited during the first week 
of June of 1980; no adults were collected and the few larvae found 
were all in the fourth and fifth stadia. 

As noted previously Schinia verna is most closely related to S. 
honesta, which is distributed in montane western North America from 
southern British Columbia southward to the Rocky Mountains of Col- 
orado and the Sierra Nevada of California. The new species is smaller 


VOLUME 37, NUMBER 1 23 


and more delicately colored than is honesta. The food plant and im- 
mature stages of honesta are unknown. It will be interesting to as- 
certain how closely the immatures resemble those of S. verna. 


ACKNOWLEDGMENTS 


I take pleasure in naming the new species after my wife, Verna, in thanks for her 
continuing assistance in the field. I also acknowledge the assistance of Mr. Eric Rock- 
burne of the Biosystematics Research Institute for the preparation of genitalic slides 
and of illustrations accompanying this paper. Mr. Wm. Shewfelt was most hospitable 
in showing me various topographic features in the region of his homestead in southern 
Manitoba. 


LITERATURE CITED 


GrROTE, A. R. 1881. New western moths. Papilio 1:75-78. 

HARDWICK, D. F. 1958. Taxonomy, life history, and habits of the elliptoid-eyed species 
of Schinia (Lepidoptera: Noctuidae) with notes on the Heliothidinae. Can. Ento- 
mol. Suppl. 6:116 pp. 


Journal of the Lepidopterists’ Society 
37(1), 1983, 24-28 


A NEW GENUS AND NEW SPECIES OF GEOMETRID 
MOTH FROM TEXAS 


DOUGLAS C. FERGUSON 


Systematic Entomology Laboratory, IIBIII, Agricultural Research Service, USDA, 
% U.S. National Museum of Natural History, Washington, D.C. 20560 


ABSTRACT. A new genus and new species of geometrid moth are described from 
West Texas. Although clearly belonging to the subfamily Ennominae, tribe Ouraptery- 
gini, this distinctive moth seems to have no close relatives. Its larva and foodplant are 
unknown. 


A geometrid from West Texas, first brought to my attention by Mr. 
André Blanchard 10 years ago, is undescribed and furthermore, does 
not fit the description of any known genus. It is clearly a member of 
the very large, possibly heterogeneous tribe Ourapterygini by reason 
of the single, massive, asymmetrical furca of the male genitalia, oc- 
curring in combination with a double accessory cell in the forewing, 
and it shows some affinity with the group that includes such genera 
as Lychnosea Grote, Caripeta Walker, Snowia Neumogen, Destutia 
Grossbeck, Besma Capps, Lambdina Capps, Cingilia Walker, Nepy- 
tia Hulst, Eusarca Hubner, and Somatolophia Hulst. The Texas 
species does not fit any of these, however, nor any of the neotropical 
genera that were investigated in the effort to place it. Species of most 
ourapterygine genera typically have a cluster or transverse row of 
hooklike spines at the extremity of the gnathos. The present species 
lacks these but instead has an extended, rather massive and recurved 
gnathos unlike any other. The so-called furca is also unusually en- 
larged but shows every indication of being homologous to those of 
the genera mentioned above. 

More than anything else the moth has the appearance of a small, 
pale, tawny-yellow, evenly colored Sicya morsicaria (Hulst), or even 
a small, narrow-winged Tetracis crocallata Guenée (although the 
forewing has only the slightest suggestion of an angulate outer mar- 
gin). However, the genitalia at once remove it from close association 
with either of those genera, and indeed the species seems to have no 
really close relatives. 


Sicyopsis blanchardata, new genus, new species 


(Figs. 1-6) 


As S. blanchardata is the only included species, it is not possible 
to differentiate between generic and specific characters; thus, the de- 
scriptions are combined. 


VOLUME 37, NUMBER l AS) 


Q | ) off 
oa | 
k ] by 
Fics. 1, 2. Sicyopsis blanchardata, n. gen., n. sp.: 1, holotype male; 2, allotype 
female (about natural size). 


Zz 


Description. Front smooth, convex, covered with closely appressed, elongated, mostly 
singly notched scales, its surface in lateral profile nearly parallel to, and just slightly 
farther out than that of adjoining eye surfaces; front about 1% times as wide at top as 
at bottom; eyes of both sexes large, each about equal in width (in radial view) to length 
of labial palpi, but slightly smaller in female than in male; palpi short, those of male 
usually slightly exceeding front, of female not exceeding front; tongue well developed; 
male antennae simple but thickened dorsoventrally, compressed, with quadrate, finely 
setose segments (prismatic); female antennae simple, filiform; legs with tibiae hardly 
swollen, epiphyses extending to ends of foretibiae, and both pairs of hindtibial spurs 
present in both sexes; vestiture of thorax predominantly of long, hairlike scales, inter- 
mixed with a few short, broad scales. 

Wings somewhat narrow, forewings with costa nearly straight, outer margin slightly 
angulate in females, usually evenly convex in males. Forewings light tawny yellow to 
cream colored, with or without an almost straight, light-brown, oblique, postmedial 
band from near middle of inner margin to a point on costa just basad of apex, and a 
small, brown discal spot; fringes concolorous with wings; hindwings paler, unmarked; 
underside unmarked. Venation (Fig. 3) typical of group of genera to which species 
seems to belong; for example, almost exactly like that of Destutia excelsa (Strecker), 
except that accessory cells of forewing variably less elongated in the four specimens 
examined for venation characters. Length of forewing: holotype, 14 mm; other males, 
13-14 mm; allotype, 14 mm; other females, 13-15 mm. 

Male genitalia (Figs. 5, 6). Uncus, gnathos, and costa of valve stout, heavily sclero- 
tized; gnathos elongated and abruptly recurved at about the middle, broad and flattened 
toward end but tapering to a blunt point apically; small, lightly sclerotized socii pres- 
ent; juxta invaginated, pouchlike, with a large, broad, single, curved, asymmetrical 
furca arising from it. As in related forms, ventral lamella of furca attached to juxta, and 
dorsal lamella to a thinly sclerotized plate of the anellus; furca bearing group of short 
spines toward its tapered extremity, transtilla incomplete; aedeagus slender, with 7-8 
small cornuti. 

Female genitalia (Fig. 4). Ductus bursae and ostium forming a nearly straight, stout, 
rigidly sclerotized, funnel-shaped unit; sterigma large, doubly invaginated as shown. 
Lack of signum unusual for the tribe. 

Types. HOLOTYPE MALE (Fig. 1): Smith Canyon, 5750 ft, Guadalupe Mountains, 
Culberson Co., Texas, 22 May 1973, D. C. Ferguson. ALLOTYPE FEMALE: Panther Pass, 
6000 ft, Chisos Mountains, Brewster Co., Texas, 2 June 1973, D. C. Ferguson. PARA- 
TYPES: | 6, same data as for holotype; 1 d, 1 2, McKittrick Canyon, 5000 ft, Guadalupe 
Mountains, Culberson Co., 23 May 1973, D. C. Ferguson; 4 66, Bear Canyon, 5400 
ft, Guadalupe Mountains, 3 Sept. 1969, A. & M. E. Blanchard; 4 66, 2 22, Bear 
Canyon, 5700 ft, Guadalupe Mountains, 4 Sept. 1969, A. & M. E. Blanchard; 1 6, Pine 
Canyon, 5700 ft, Guadalupe Mountains, 28 Aug. 1967, A. & M. E. Blanchard; 9 34, 
Sierra Diablo Wildlife Management Area, 6000 ft, 20 mi. NNW of Van Horm, Culberson 
Co., 20 May 1968 (1), 5 June 1969 (4), 14 July 1971 (2), 30 Aug. 1970 (1), 24 Sept. 1967 


26 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


0 5 10 


Fics. 3, 4. Sicyopsis blanchardata: 3, wing venation; 4, female genitalia, from 
USNM slide 56097 (Guadalupe Mts.). 


(1), A. & M. E. Blanchard; 1 36, same locality, 30 May 1973, D. C. Ferguson; 10 66, 
same locality, 30 Aug. 1970 (6), 31 Aug. 1970 (4), J. G. Franclemont; 2 2 2, same locality 
and collector, 30 Aug. 1970; 1 2, 10 mi. north of Van Horn, Culberson Co., 2 Sept. 
1979, E. C. Knudson; 23 66, Green Gulch, 5500 ft, Chisos Mountains, Big Bend 
National Park, Brewster Co., 10 May 1966 (1), 14 May 1966 (1), 27 June 1965 (2), 1 
July 1965 (1), 14 Sept. 1971 (5), 2 Oct. 1967 (10), 5 Oct. 1965 (2), 7 Oct. 1965 (1), A. & 
M. E. Blanchard; 1 ¢, same locality and collectors, 27 Aug. 1965; 1 2, same locality, 
6 June 1973, D. C. Ferguson; 10 66, Chisos Basin, Big Bend National Park, 29 June 
1965 (1), 30 Aug. 1964 (1), 27 Sept. 1965 (4), 2 Oct. 1966 (3), 7 Oct. 1966 (1), A. & M. 
E. Blanchard; 3 22, same locality and collectors, 30 Aug. 1964, 24 Sept. 1963, 5 Oct. 
1967; 1 5, same locality, 14 May 1977, E. C. Knudson; 3 66, Government Spring, Big 
Bend National Park, 13 May 1966, 29 Sept. 1965, 4 Oct. 1967, A. & M. E. Blanchard; 
9 $6, Oak Spring, Big Bend National Park, 4 Oct. 1965 (2), 5 Oct. 1967 (5), 5 Oct. 1965 
(2), A. & M. E. Blanchard; 1 ¢, K-Bar Research Station, Big Bend National Park, 25 
Sept. 1971, A. & M. E. Blanchard. 

All localities cited are in West Texas. The Guadalupe Mountain localities are in 
Guadalupe Mountains National Park in canyons on the east side of the range. The 
holotype, allotype, and some of the paratypes are in the collection of the U.S. National 
Museum of Natural History; other type material is in the collections of A. Blanchard, 
J. G. Franclemont, and E. C. Knudson. 


VOLUME 37, NUMBER l Di 


1 mm 


Fics. 5, 6. Sicyopsis blanchardata, male genitalia: 5, ventral view, aedeagus re- 
moved, from slide AB 1257 (Big Bend National Park). 6, ventrolateral view, right valve 
removed, aedeagus in situ, from slide AB 1376 (Green Gulch). Drawings by A. Blan- 
chard and the author. 


28 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Distribution. All known localities are given above. Not known from Mexico, but 
inasmuch as many were collected virtually within sight of the border, it undoubtedly 
occurs there. . 

Flight period. 10 May-7 October. 

Early stages. Unknown. 

Remarks. This species has been named in honor of Mr. André Blanchard of Houston, 
Texas, who collected most of the material and who personally conducted me to the 
localities where those that I collected were found. He also provided the drawings for 
Figs. 3, 5, and 6, which I modified for publication. I also acknowledge the assistance 
of A. Blanchard, E. C. Knudson, and J. G. Franclemont for providing information or 
access to their collections. 


Journal of the Lepidopterists’ Society 
37(1), 1983, 28 


BOOK REVIEW 


A REVISION OF PHYCIODES HUBNER AND RELATED GENERA, WITH A REVIEW OF THE 
CLASSIFICATION OF THE MELITAEINAE (LEPIDOPTERA: NYMPHALIDAE) by L. G. Hig- 
gins. 29 October 1981. Bulletin of the British Museum (Natural History), Entomological 
Series, Vol. 43, No. 3, pp. 77-243, 490 figs. 


This concludes Higgins’ 30-year study of the melitaeines of the world. In the present 
volume the Phyciodini and ten genera are described and named for the first time. At 
the species-group level eight new names are introduced. Many shifts from the species- 
group arrangements of Arthur Hall (1928-1930) and William T. M. Forbes (1945, 1946) 
have been necessary. Higgins’ work had the advantage of the huge collections of the 
British Museum (Nat. Hist.) and Hall’s very large collection in the Booth Museum, 
Bristol, England. In the British Museum alone Higgins had before him 76% of the 
types involved. 

For each name noted in his complete synonymies for every accepted species, Higgins 
notes the author, original citation, type locality and repository of the type. In the cases 
of five authors the type repositories evaded him. Reakirt’s types are in the Strecker 
Collection, Field Museum of Natural History, currently on long-term loan to the Allyn 
Museum in Sarasota, Florida. David Bauer’s types are in his own collection, South 
Lake Tahoe, California. Brehme’s types may be with the material from the Brooklyn 
Museum now in the U.S. National Museum of Natural History, Washington, D.C. I 
have no idea where Fender's type of P. mylitta ab. ““maceyi’ may be. Cockerell’s two 
abs. of camillus, “rohweri” and “tristis,” probably have been lost. 

None of the newly proposed generic names affect species names in the United States 
and Canada. A comparison between Higgins’ arrangement and Brown’s arrangement 
of Phyciodes in the new Catalogue/Checklist shows the usual differences in assignment 
of status, i.e., species, subspecies and synonyms, between two serious students of any 
topic. Higgins’ approach is more conservative than Brown’s. There are two differences 
that are more substantial: Where Higgins used campestris Behr, 1863 (following Barnes 
& McDunnough, 1917), Brown reverted to the earlier use of pratensis Behr, 1863 as 
the species name, following Strecker’s Catalogue of 1876 as a “first revision.’ Whereas, 
Higgins overlooked callina Boisduval, 1869, as a misspelling of collina Behr, 1863, 
Brown considers it a subspecies of mylitta, with arizonensis Bauer being a synonym. 

No price is noted on my copy of the revision. I am sure that Classey will be handling 
it and suggest that anyone interested write to him at his England headquarters. 


F. MARTIN BROWN, Wright-Ingraham Institute, Colorado Springs, Colorado 80904. 


Journal of the Lepidopterists’ Society 
37(1), 1983, 29-37 


PROLONGED DIAPAUSE AND PUPAL SURVIVAL 
OF PAPILIO ZELICAON LUCAS 
(LEPIDOPTERA: PAPILIONIDAE) 


STEVEN R. Sims! 


Department of Entomology, 840 Method Road, Unit I, 
North Carolina State University, Raleigh, North Carolina 27650 


ABSTRACT. Pupae of Papilio zelicaon are capable of at least a two-year diapause. 
Univoltine populations utilizing ephemeral native host plants had a greater incidence 
of prolonged diapause than multivoltine populations using the introduced Foeniculum 
vulgare. A laboratory strain, derived from multivoltine individuals and selected for 
decreased diapause, exhibited a corresponding loss of prolonged diapause. Mortality 
was high among pupae overwintering twice and complete in three-year-old individuals. 
Pupal water and dry material (fat body, etc.) weight loss was gradual over a 541 day 
period and paralleled survivorship. Prolonged diapause is regarded as an adaptive 
response to environmental uncertainty, especially that related to the phenology of host 
plants associated with univoltine and multivoltine populations. 


Univoltine (one brood/year) populations of the swallowtail butter- 
fly, Papilio zelicaon Lucas, typically occur in rocky or exposed areas 
at mid to high elevations in the California Coast Range and Sierra 
Nevada. These populations feed on native species of Umbelliferae 
(especially Angelica, Lomatium, Pteryxia, and Tauschia spp.) that are 
suitable for larval development approximately 2 to 4 months each year 
(Emmel & Shields, 1978). Multivoltine (several broods/year) popu- 
lations occur in California coastal, Sacramento, and San Joaquin Val- 
ley areas, where they feed primarily on the introduced weed Foeni- 
culum vulgare Mill. (Shapiro, 1974a, b) but occasionally attack Citrus 
spp. (Coolidge, 1910; Horton, 1922; Shapiro & Masuda, 1980). The 
temporal suitability of Foeniculum and Citrus as larval hosts varies 
among individual plants and location but, based on the annual avail- 
ability of new foliage and succulent leaves, host suitability ranges 
from 8 to 12 months (Emmel & Shields, op. cit.). 

P. zelicaon undergoes a photoperiod-induced pupal diapause, the 
incidence of which is modified by temperature and host plant species 
(Sims, 1980). The intensity of pupal diapause (=duration under spec- 
ified environmental conditions) varies between individuals and among 
populations; individuals from a lab strain have reduced response to 
diapause-inducing conditions (Sims, in press). Some southern Califor- 
nia populations (Oliver, 1969) appear to have reduced chilling re- 
quirements for diapause termination compared to central California 
populations. Some pupae remain in a diapause of at least two years’ 


' Current address: University of Florida AREC, 18905 SW 280th Street, Homestead, Florida 33031. 


30 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


duration, either under artificial indoor conditions (Hynes, 1949) or 
outdoors exposed to naturally varying temperature and photoperiod 
(Emmel & Shields, op. cit.; Emmel, pers. comm.). The purpose of this 
study was to document the occurrence of prolonged diapause in sev- 
eral univoltine and multivoltine P. zelicaon populations and to obtain 
estimates of overwintering survival in normal and prolonged diapause 
pupae. Observations were made on progressive pupal weight loss as 
one possible mortality agent of diapausing pupae. 


METHODS 


Ova were obtained from females field-collected in univoltine (Butts 
Canyon, Napa Co., CA, el. 500 m, and Donner Pass, Placer Co., CA, 
el. 2100 m) and multivoltine (Suisun Marsh, Solano Co., CA, el. <10 
m and Rancho Cordova, Sacramento Co., CA, el. <30 m) populations. 
Techniques used to establish and maintain colonies were similar to 
those of Sims (1980). Larvae were reared on Foeniculum under a 14 
h light/10 h dark (LD 14:10) photoperiod at 23.5°C. Diapause pupae 
reared under similar conditions were also obtained from the F3—F; 
generations of a laboratory stock which originated from the two mul- 
tivoltine populations. Within 90 days of pupation, diapause pupae (in 
1 pt. cardboard containers) were placed in a sheltered outdoor cage 
exposed to normal fluctuations of temperature, humidity, and photo- 
period at Davis, Yolo Co., CA, el. 10 m, during the years 1975-1977. 
Emergence and mortality data were recorded daily during spring and 
summer and weekly during the remainder of the year. 

Desiccation tolerance of diapausing pupae was studied by record- 
ing survivorship and wet weights of 24 pupae from Donner Pass over 
a 541 day period. Pupae were maintained at LD 15:9, 26.5°C with a 
relative humidity of 35 + 5%. The laboratory conditions used sub- 
jected the pupae to much more desiccation stress than they are likely 
to experience in their normal habitat. Mortality in pupae was indi- 
cated by greatly accelerated water loss. The presence of unidentified 
disease in many dead pupae was suggested by abdominal swelling 
and exposure of the intersegmental membrane revealing discol- 
ored internal body fluids. Water and dry weight loss over the 541 days 
was estimated by weighing samples of pupae at the beginning and 
end of this period. Weighed living pupae were slit open, dried at 
90°C for 48 h in a vacuum oven, and reweighed. 


RESULTS 


Progeny of four females from Rancho Cordova (combined n = 69) 
and three females from Suisun Marsh (n = 64) were compared to prog- 


VOLUME 37, NUMBER | ol 


TABLE 1. Numbers of P. zelicaon pupae diapausing a second winter inside a field 
cage in Davis, CA. 


Lab strain Multivoltine Univoltine 
# dia- # dia- # dia- 
Popula- pause/ Popula- pause/ Popula- pause/ 
tion/ total % dia- tion/ total % dia- tion/ total % dia- 
female alive pause female alive pause female : alive pause 
F,/1 0/8 0.0 Rancho LEW 7 68.4 Donner/1 4/4 100.0 
Cordova/1 
F,/2 0/8 0.0 Rancho 4/16 25.0 Donner/2 8/12 66.7 
Cordova/2 
F,/3 C/A? 0.0 Rancho 1/28 3.6 Donner/3 1/3 Soro 
Cordova/3 
F,/4 0/8 0.0 Rancho 0/8 0.0 Butts 6/13 46.1 
Cordova/4 Canyon/1 
F,/1 OA? 0:0 Suisun/1 5/9 50.6 
F,/2 0/5 0.0 Suisun/2 A/17 DABS, 
Suisun/3 0/38 0.0 
Total? 0/52 0.0 Total DAS LES" Ko Ko: Total 19/32 59.4 


4 Univoltine total is significantly greater than laboratory or multivoltine (Duncan’s Multiple Range Test, P < 
0.05). 


eny of three Donner Pass and one Butts Canyon female (n = 32). Six 
females from the multivoltine-derived laboratory colony provided 52 
pupae for comparison with the wild type populations. Results are 
given in Table 1..The proportion of living non-emerged pupae of 
those overwintering for the first year was arc-sine transformed using 
the tables provided by Mosteller and Youtz (1961). Transformed data 
were analyzed using ANOVA procedures (Sokal & Rohlf, 1969) and 
Duncan's Multiple Range Test (DMR). Differences among the pop- 
ulations within the uni- and multivoltine areas and between the F; 
and F, generations of the laboratory strain were not significant; there- 
fore, a comparison was made between the combined laboratory strain, 
multivoltine, and univoltine populations. The difference between 
these groups was significant (F,,, = 10.26, P < 0.002). No pupae from 
the laboratory strain overwintered a second time; whereas, 19% of the 
multivoltine and 59% of the univoltine individuals did so. The dif- 
ference between the latter two groups is significant (P < 0.05, DMR). 

Emergence and survivorship of diapausing pupae older than one 
year was examined in 97 laboratory strain (generations F;—F;) indi- 
viduals that had been previously chilled from November 1974 to March 
foo tor 126 days at 2°,\5°, 11°, or 16°C at LD 0:24. Of 97 pupae 
maintained in an outdoor cage over the 1975-1976 and 1976-1977 
winters, 24 66 and 27 2 ° developed the first spring. Fifteen of these 
adults were either dead inside the pupa or emerged very weak and/ 


OZ JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


100 
90 
80 
70 


60 


PERCENT ALIVE 


50 


PERCENT INITIAL WEIGHT 


40 


30 


(0) 
(0) 80 160 240 320 400 480 560 


DAYS 


Fic. 1. Weight loss (percent of initial weight remaining) and survivorship of 24 
Papilio zelicaon pupae under LD 15:9, 26.5°C conditions. 


or crippled. Thus, only 37.1% (36/97) of the pupae produced adults 
that appeared to have adequate vigor for normal reproductive activity. 
The 44 unemerged pupae (45.6%) died without showing signs of de- 
velopment, while 2/97 (2.1%) remained alive in diapause. These latter 
two pupae plus 13 surviving two-year-old uni- and multivoltine pupae 
died while overwintering a third time. There was no difference be- 
tween percent emergence or mortality of individuals previously chilled 
at the different temperatures. Prolonged diapause in the populations 
of P. zelicaon studied, therefore, appears to be normally of two years’ 
maximum duration. Mortality is high in second-year pupae and com- 
plete in three-year-old individuals. 

Figure | illustrates weight loss and survivorship of 24 P. zelicaon 
pupae over a 541 day period. A steep decline in weight, correspond- 
ing to water loss during hardening of the pupal integument, occurred 
during the first week of pupation. Subsequent weight loss was gradual 
and nearly linear over time. The survivorship curve in Figure | rough- 
ly parallels the progressive loss of pupal weight. The highest mortality 
rate (50%) occurred during the first 120 days; the remaining pupae 
had a high probability of survival (83%) to day 360. Although weight 


VOLUME 37, NUMBER 1 33 


TABLE 2. Depletion of water and dry weight of P. zelicaon pupae over a 541 day 
period at LD 15:9, 26.5°C conditions and RH = 35 + 5%. 


Pupal Total Percent Total 


age Total water dry weight dry weight Percent percent 
(days)? (g) (g) Water/dry weight loss water loss weight loss 
1 (10) 0.814 0.238 3.419 — _— — 
541 (6) 0.593 0.160 3.700 32.68 2 NA 28.39 


a Number in parentheses indicates numbers of pupae sacrificed. 


loss and survivorship are probably not related in a simple way, it is 
possible that mortality of some pupae, especially those older than one 
year, occurred due to water or energy reserve depletion. The six sur- 
viving pupae at day 541 were 84% (x day 1 = 0.955 g; x day 541 = 
0.802 g) of their own day 1 weight. 

Table 2 provides an analysis of water and dry weight loss over 541 
days. Rates of weight loss of both water (27.1%) and dry material (fat 
body, etc.) (32.7%) are similar, although the actual weights of water 
(0.221 g) and dry matter (0.078 g) lost differ greatly. Similar initial 
and final ratios of water to dry weight (3.419 to 3.700) suggest ho- 
meostatic maintenance of water/fat body quantities by the pupae. 


DISCUSSION 


The data presented here provides evidence for both individual fe- 
male brood variation and population differentiation in the incidence 
of prolonged pupal diapause in P. zelicaon. Prolonged diapause was 
most frequent in univoltine populations and least frequent in a lab- 
oratory strain founded from multivoltine individuals. 

The laboratory strain of P. zelicaon was selected for continuous 
development at LD 14:10, 23°C; thus, pupae diapausing under these 
“long-day” conditions were eliminated from the culture. By the F, 
generation this procedure had significantly decreased the incidence 
of diapause (Sims, in press) as well as eliminated any manifes- 
tation of prolonged diapause (Table 1). This dual response to selection 
suggests that the regulation of both diapause initiation and diapause 
intensity is under genetic control. 

Comparison in Davis of diapause intensity in P. zelicaon from low 
elevation (up to 500 m) coast range, coastal, and central valley pop- 
ulations seems justified by the similarity of winter temperature monthly 
means (8—11°C) in these areas. Higher elevation (750-2100 m) Sierra 
Nevada populations experience much lower winter temperature 
means, often spending several months under snow cover with tem- 
peratures at or near 0°C. Despite this, pupae from populations at all 
elevations examined exhibit a maximum diapause termination re- 


34 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


sponse (under constant temperatures) at 11°C (Sims, 1980). Thus, 
overwintering of higher elevation pupae under lower elevation valley 
conditions should provide an adequate approximation of prolonged 
diapause frequency. 

Prolonged diapause has been found in a number of other temperate 
zone Lepidoptera that overwinter as pupae (Biliotti, 1953; Rabb, 1966; 
Powell, 1974; Prentiss, 1976; Shapiro, 1980) although it is not limited 
to the pupal stage (Yothers & Carlson, 1941; Surgeoner & Wallner, 
1978). 

Differences in photoperiod response and diapause intensity of lat- 
itudinally and altitudinally separated populations are commonly ob- 
served (Danilevskii, 1965; Beck, 1980; Sims, 1980) but interpopu- 
lation differentiation and temporal variability in the occurrence of 
prolonged diapause is less well known. Significant interpopulation 
variation in prolonged diapause has been found in sawflies (Prebble, 
1941; Eichhorn, 1977), Cecidomyiidae (Sunose, 1978) and Lepidop- 
tera (Surgeoner & Wallner, op. cit.). Prebble (op. cit.) obtained results 
similar to those of this study; uni- and bivoltine populations of Gil- 
pinia polytoma (Hartig) (Hymenoptera, Diprionidae) were found to 
have considerably more prolonged diapause than multivoltine pop- 
ulations. 

A major ecological difference between univoltine and multivoltine 
P. zelicaon populations is the growth phenology of the host plants 
(Emmel & Shields, op. cit.). Native perennial species of umbellifers 
used by lower elevation univoltine populations senesce rapidly with 
the advent of warm, dry weather in May and June. Pteryxia terebin- 
thina (Hook.) C. & R., a host used by the Donner Pass and other high 
elevation Sierra Nevada populations (Emmel & Emmel, 1974), has a 
similar senescence in mid-July to August, placing a severe food lim- 
itation on occasional partial second broods unable to find or utilize 
alternate host species (Sims, 1980). Possibility of killing frosts in 
early fall increases the hazards for second brood individuals at higher 
elevations. The unpredictable quality and quantity of available food 
plants for larval development during the latter part of the growth 
season presents a considerable environmental risk for P. zelicaon. 
This food plant risk is partially buffered on Foeniculum since this 
“weedy species has a longer growth season and occurs in a greater 
diversity of habitats than native hosts. In mesic areas Foeniculum may 
have tender young foliage throughout the season, and the plant com- 
monly sends out fresh growth following disturbance. Variation in pre- 
cipitation as it effects host plant growth is not the only possible en- 
vironmental risk that may influence prolonged diapause in P. zelicaon. 


VOLUME 37, NUMBER 1 35 


Studies on other insects have demonstrated the adaptive value of pro- 
longed diapause in the avoidance of biotic hazards such as disease, 
predators, and parasites (Price & Tripp, 1972; Minder, 1973). 

Relatively dry areas of the temperate zone, including the Mediter- 
ranean climate of central California, tend to have particularly variable 
rainfall patterns (Rumney, 1969). Records from Sacramento provide a 
good example of this variability. Here, 120 years of data (1849-1969) 
show a mean seasonal rainfall of 45.5 cm with a range of 11.9-92.5 
cm (Figgins, 1971). In seven of the 120 years the seasonal total was 
less than one-half the mean and the 1975-1976 and 1976-1977 seasons 
subjected populations to consecutive extreme drought years (15.6 and 
19.5 cm rainfall, respectively). The effect of drought on host plants 
and population levels and/or population persistence through time is 
still poorly understood for P. zelicaon. For Euphydryas editha Bois- 
duval, a species frequently co-occurring with P. zelicaon in central 
California, drought can reduce population size through larval starva- 
tion (White, 1974) or be a factor involved in local population extinc- 
tion (Ehrlich et al., 1980). 

Within the context of current thought on life history adaptations in 
insects, the presence of prolonged diapause in P. zelicaon represents 
a ‘bet-hedging”’ strategy in which individual females spread the risk 
of their reproductive effort over more than one season (den Boer, 
1968; Stearns, 1976). This explanation implies, but does not demon- 
strate, the unpredictable risk of complete reproductive failure among 
the diapause progeny from a single female emerging in a given year. 
It is also possible that, even without the risk of a catastrophic season, 
the value of R, (net reproductive or replacement rate during a growth 
season) will be small enough in some years to select for the allocation 
of progeny into both normal and extended diapause categories (Istock, 
1981). On an evolutionary scale the presence of prolonged diapause 
suggests that the long-term benefits of this response outweigh the 
disadvantages of the increased mortality rate demonstrated in this 
study. 


ACKNOWLEDGMENTS 


J. F. Emmel, S. O. Mattoon, and H. A. Tyler provided living material and biological 
information. Drs. C. L. Judson and R. W. Thorp assisted on an earlier draft, and C. 
Satterwhite aided in manuscript preparation. Dr. A. M. Shapiro was a source of intel- 
lectual stimulation and encouragement throughout this study. 


LITERATURE CITED 


BEcK, S. D. 1980. Insect Photoperiodism. Second Edition. Academic Press, New York. 
387 pp. 


36 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


BILIoTTI, E. 1953. Importance et signification des arréts de dévelopement au stade 
nymphal chez Thaumetopea processionea L. C. R. Acad. Sci. 236:1703-1705. 
BoER, P. J. DEN. 1968. Spreading of risk and stabilization of animal numbers. Acta 

Biotheor. 18:165—-194. 

CooLipGE, K. R. 1910. A California orange dog. Pomona Coll. J. Entomol. 2:333- 
334. 

DANILEVSKU, A. S. 1965. Photoperiodism and Seasonal Development of Insects. Oliver 
and Boyd, London. 283 pp. 

EHRLICH, P. R., D. D. MURPHY, M. C. SINGER, C. B. SHERWOOD, R. R. WHITE & I. L. 
BROWN. 1980. Extinction, reduction and stability: the responses of checkerspot 
butterflies to the California drought. Oecologia (Berl.) 46:101—105. 

EICHHORN, O. 1977. Studies on the autecology of the common pine sawfly Diprion 
pini (L.). I. Strain-specific differences in emergence and diapause. Z. Angew. Ento- 
mol. 82:395-414. 

EMMEL, J. F. & O. SHIELDS. 1978 (1979). Larval food plant records for Papilio zeli- 
caon in the western United States and further evidence for the conspecificity of 
P. zelicaon and P. gothica. J. Res. Lepid. 17:56—67. 

EXMMEL, T. C. & J. F. EMMEL. 1974. Ecological studies of Rhopalocera in a Sierra 
Nevadan community—Donner Pass, California. V. Faunal additions and foodplant 
records since 1962. J. Lepid. Soc. 28:344-348. 

FIGGINS, W. E. 1971. Climate of Sacramento, California. NOAA Tech. Mem. NWS- 
WR65. 63 pp. 

HORTON, J. R. 1922. A swallowtail butterfly injurious to California orange trees. Mon. 
Bull. Calif. State Hortic. Comm. 11:377-387. 

Hynes, V. P. 1949. Papilio pupa emerging after two winters. Lepid. News 3:64. 

Istock, C. A. 1981. Natural selection and life history variation: theory plus lessons 
from a mosquito. Pp. 113-126, in R. F. Denno & H. Dingle, (eds.). Insect Life 
History Patterns: Habitat and Geographic Variation. Springer-Verlag, New York. 

MINDER, I. F. 1973. Prolonged diapause in larvae of Neodiprion sertifer. Zool. Zh. 
52:1661-1670. 

MOSTELLER, F. & C. YouTz. 1961. Tables of the Freeman-Tukey transformations for 
the binomial and Poisson distributions. Biometrika 48:433-440. 

OLIVER, C. G. 1969. Experiments on the diapause dynamics of Papilio polyxenes. J. 
Insect Physiol. 15: 1579-1589. 

POWELL, J. A. 1974. Occurrence of prolonged diapause in ethmiid moths. Pan-Pac. 
Entomol. 50:220-225. 

PREBBLE, M. L. 1941. The diapause and related phenomena in Gilpinia polytoma 
(Hartig). V. Diapause in relation to epidemiology. Can. J. Res., D 19:437-454. 
PRENTISS, J. B. 1976. Time variations of pupal stage of Eupackardia calleta (Satur- 

niidae). J. Lepid. Soc. 30:187. 

Prick, P. W. & H. A. Tripp. 1972. Activity patterns of parasitoids on the Swaine jack 
pine sawfly, Neodiprion swainei (Hymenoptera: Diprionidae), and parasitoid im- 
pact on the host. Can. Entomol. 104:1003-1016. 

RABB, R. L. 1966. Diapause in Protoparce sexta (Lepidoptera: Sphingidae). Ann. 
Entomol. Soc. Amer. 59:160-165. 

RUMNEY, G. R. 1969. Climatology and the World’s Climates. Macmillan, London. 
656 pp. 

SHAPIRO, A. M. 1974a. Butterflies of the Suisun Marsh, California. J. Res. Lepid. 13: 
191-206. 

1974b. The butterfly fauna of the Sacramento Valley, California, J. Res. Lepid. 

13:73-82, 115-122, 137-148. 

1980. Egg-load assessment and carryover diapause in Anthocharis (Pieridae). 

|. Kepid. Soc, 34:307-315:. 

& K. K. MAsupDA. 1980. The opportunistic origin of a new citrus pest. Calif. 
Agr. 34:4-5. 

SIMS, S. R. 1980. Diapause dynamics and host plant suitability of Papilio zelicaon 
(Lepidoptera: Papilionidae). Amer. Midl. Nat. 103:375-384. 


VOLUME 37, NUMBER l Sr 


SOKAL, R. R. & F. J. ROHLF. 1969. Biometry. W. H. Freeman & Co., San Francisco, 
CAG (Gipp: 

STEARNS, S.C. 1976. Life-history tactics: a review of the ideas. Quart. Rev. Biol. 51: 
3-47. 

SUNOSE, T. 1978. Studies on extended diapause in Hasegawaia sasacola Monzen 
(Diptera, Cecidomyiidae) and its parasites. Kontya 46:400-415. 

SURGEONER, G. A. & W. E. WALLNER. 1978. Evidence of prolonged diapause in 
prepupae of the variable oakleaf caterpillar, Heterocampa manteo. Environ. Ento- 
mol. 7: 186-188. 

WHITE, R. R. 1974. Food plant defoliation and larval starvation of Euphydryas editha. 
Oecologia (Berl.) 14:307-315. 

YOTHERS, M. A. & F. W. CARLSON. 1941. Orchard observations of the emergence of 
codling moths from two-year-old larvae. J. Econ. Entomol. 34:109-110. 


Journal of the Lepidopterists’ Society 
37(1), 1983, 37 


OBITUARY 
Dr. RENE Licuy (1896-1981) 


Professor Lichy was an outstanding lepidopterist who lived and worked for many 
years in Venezuela. I first knew him in 1938, and we have exchanged papers and 
occasional letters ever since. He taught on the faculty of the Universidad Central de 
Venezuela, was a member of the Academia de Ciencias Fisicas-matematicas y Naturales 
de Venezuela, an honorary member of both the Sociedad Venezolana de Ciencias 
Naturales and Sociedad Venezolana de Entomologia. He was the honorary curator for 
Lepidoptera at the Museo de Ciencias Naturales in Caracas. He joined the Lepidop- 
terists’ Society soon after its foundation. 

The last twenty years or so of his life were spent in France, the land of his origin. 
There he was associated with the Department of Entomology at the Museum d Histoire 
Naturelle in Paris. He continued his great interest in Venezuelan Lepidoptera and 
furthered his world-wide studies of Sphingidae. René Lichy died on 6 April 1981. He 
is survived by three children and three grandchildren. His daughter and her three 
children live in France. Also, refer to Freiche & Lemaire (1981, Bull. Soc. Ent. France 
86(9&10):313-314). 


F. MARTIN BROWN, 6715 S. Marksheffel Rd., Colorado Springs, Colorado 80911. 


Journal of the Lepidopterists’ Society 
37(1), 1983, 38-45 


NOTES ON THE BIOLOGY OF 
AGONOPTERIX ALSTROEMERIANA (CLERCK), 
WITH DESCRIPTIONS OF THE IMMATURE 
STAGES (OECOPHORIDAE) 


M. BERENBAUM! 


Department of Entomology, 135 Insectary, Cornell University, 
Ithaca, New York 14853 


AND 


S. PASSOA 
219-66 67th Ave., Bayside, New York 11364 


ABSTRACT. Larvae collected from Conium maculatum L. (Umbelliferae) in 
Tompkins County, New York, were reared and determined as Agonopterix alstroe- 
meriana (Clerck), recorded in the United States only since 1973. The larva forms a 
tubular roll on leaves of Conium maculatum, its sole host. Pupation occurs in late May 
and early June; adults emerge in mid- to late June. 

The last instar larva is greenish in color, with a dorsal and two subdorsal dark green 
stripes. Earlier instar larvae are predominantly yellow with black head capsules. UI- 
timate instar larvae of A. alstroemeriana can be distinguished from other species of 
Agonopterix by the black posterior margins of the epicrania, the conspicuous pinacula, 
and scalelike setae dorsal to the tarsal claws. 


Larvae of Agonopterix alstroemeriana (Clerck) were collected and 
reared on Conium maculatum L. (Umbelliferae) at three different 
sites in Tompkins County, New York, during the summers of 1977, 
1978, and 1979. A. alstroemeriana is recorded in North America only 
since 1973 (Hodges et al., in press); since it is not included in the 
recent revision of the Oecophoridae of North America (Hodges, 1974), 
this paper documents for the first time the biology of this European 
species in eastern North America. 


METHODS 


Larvae were collected from C. maculatum growing in Coy Glen, 
Brooktondale, and the city of Ithaca, all in Tompkins County; C. mac- 
ulatum was one of 12 species of Umbelliferae examined at weekly 
intervals from May through August 1979 for a study on insect asso- 
ciates of umbellifers (Berenbaum, 1981). All larvae were reared on C. 
maculatum in a controlled-environment chamber, with day/night tem- 
peratures of 26.5°/15.5°C and photoperiod of 16/8 h. 

For morphological examination larvae were fixed in boiling water 
and preserved in 70% ethanol. Twenty-six reared adults and six larvae 


1 Current address: Department of Entomology, 320 Morrill Hall, University of Illinois, 505 S. Goodwin, Urbana, 
Illinois 61801. 


VOLUME 37, NUMBER l 39 


wy 


a je * at a " : 7 “ bigicte» 
} hie i y a 4 ; : 4 ae oh 
, ‘=e F A E Ag) 
BS , . mat Ry. nis ; F , 
. bey 1 
. : Pi naee J Pres 
\“ . . > - 7 . . ie A 
q * j ies ? a ess. sae > 
aes d : : ; ; Pe ~h 
es Pe, ie = y : . ? . . 
dle ae »~ wi ae: ; gt SS te ae ey M ~ eI | 


te: 


Fic. 1. Leaf rolls on Conium maculatum used as larval shelter for A. alstroemeri- 
ana. 


were examined in detail; microscopic slides of the genitalia, wings 
and larval mouthparts were stained in mercurochrome and mounted 
in balsam after clearing in clove oil. 


Natural History 


Laboratory rearing suggests that adults overwinter and lay eggs in 
early spring; this is not inconsistent with other species in the genus 
Agonopterix (Hodges, 1974). Early instar larvae can be found in 
Tompkins County in early May, at which time they form characteristic 
tight tubular leaf rolls on leaves of Conium maculatum. Weekly cen- 
sus of stands of C. maculatum revealed a steady decline in numbers 
of caterpillars during June and July, when adult emergence takes 
place (Table 1). Both flowering and nonflowering individuals of C. 
maculatum are attacked. In addition to rolling leaves, A. alstroemeri- 
ana also webs together flowers and developing seeds. Inspection of 
C. maculatum throughout July and August failed to reveal larvae; 
presumably, A. alstroemeriana is univoltine in New York, as are the 
other umbellifer-feeding oecophorids in the area (Hodges, 1974). Lar- 
vae have a tendency to abandon leaf rolls frequently, particularly after 
disturbance; on a given plant, then, a high proportion of leaf rolls are 


40 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Fics. 2, 3. Larva of A. alstroemeriana: 2, mature larva showing dorsal setae (D1, 
D2), lateral setae (L1, L2, L3), subdorsal setae (SD1, SD2), spiracle (SP), subventral 
setae (SV1, SV2), and cervical shield setae (KD1, XD2); 3, penultimate instar larva. 


vacant. Population densities can reach high levels, with up to 40 cat- 
erpillars per stem. 

A weekly census of insects on 11 other species of Umbelliferae in 
Tompkins County, some of which grow sympatrically with C. macu- 
latum (e.g., Pastinaca sativa L. and Daucus carota L.) in abandoned 
fields and ditches, did not reveal A. alstroemeriana. In New York as 
in Europe (Stainton, 1861; Schutze, 1931; Toll, 1964), C. maculatum 
appears to be the sole host for A. alstroemeriana. 


Description of Last-Instar Larva 
(terminology after MacKay, 1972) 


General. Mature larva (Fig. 2) light green with three dark green longitudinal stripes. 
Prothoracic shield concolorous with body except for black posterior margin of epicrani- 
um and small ocular spot. Fully grown larva approximately 10 mm long. Smaller larvae 
lacking green stripes, with both head and cervical shield blackish-brown (Fig. 3). 

Head (Figs. 4-9). Mandible with seven apical teeth and two lateral setae (Fig. 4). 
Molar grooves on teeth 1, 3 and 4 but most prominent on first tooth. Epicranium with 
black posterior margin (Fig. 5). Proximomedial area of hypopharyngeal complex (Fig. 
6—terminology after Godfrey, 1972) covered by very fine setae. One pair of stipular 
setae present. Spinneret (Fig. 7) with rounded apex and silk pore. Labial palpi with 
second segment reduced and third segment hairlike, with hairlike papilla on the basal 
segment. Adfrontal area and labrum as shown (Fig. 8) (terminology after Hinton, 1946). 
Adf, between epicranial and adfrontal sutures; Adf, more widely separated. Frontal 


VOLUME 37, NUMBER 1 4] 


TABLE |. Per stem number of A. alstroemeriana larvae on Conium maculatum in 
Coy Glen, Ithaca, New York (summer 1979). 


Date Mean number + S.E. 

24 May 21.4 + 5.87 
31 May 13.8 + 2.96 

7 June 11.8 + 3.20 
14 June 14.6 + 5.80 
21 June 5.8 + 1.50 
28 June 0.4 + 0.40 

5 July 0.2 + 0.20 
12 July 0.2 + 0.20 
19 July 0.0 + 0.00 


1 Five stems examined on each sampling date. 
* NoA. alstroemeriana present on subsequent sampling dates (through 31 August). 


setae located above frontal punctures; clypeal setae near the edge or outside of frontal 
triangle. Labrum with three lateral setae and three medial setae—L1 and L2 closer 
together than to L3, and M1, M2 and M3 in a triangle. On epicrania (Fig. 9), P2 only 
slightly higher than Pl and Al and A3 much longer than A2; L1 as shown. 

Thorax. On the prothorax (Fig. 2), XD1, XD2 and SD1 in a straight line; SD2 sep- 
arate from D1 and D2. Three prespiracular setae and two setae in the SV group; cervical 
shield unmarked. Chaetotaxy of metathorax: D1 and D2 closer to each other than on 
abdominal segments; SD1 and SD2 on the same pinaculum as L1 and L2; L3 widely 
separated. SV1 behind coxa. All thoracic legs with two scalelike setae dorsad of tarsal 
claw. 

Abdomen. One ventral seta present on each side of midline of abdominal segments 
(Fig. 10). D1 and D2 widely spaced; SD1 above spiracle, L1 and L2 below it and close 
to each other. L3 close to SV group. SV setae for abdominal segments 1, 2, 7, 8 and 9 
are 2, 3, 2, 1 and | respectively. Chaetotaxy of abdominal segment 9 as shown. 

Pupa. Pupa (Figs. 11-13) red-brown and pubescent, covered with dense fine hairs. 
Antennae long and curled around wing pads; vertex rounded and prominent. Meso- 
thoracic and metathoracic legs exposed; prothoracic femora concealed by maxillae. 
Segments 5 and 6 with proleg scars. Cremaster present as small tubercle with 4 hooked 
setae. Another pair of setae visible in dorsal view on each side of cremaster. Sexes 
distinguished by the position of genital openings, with ostium ductus ejacularis on the 
ninth segment and ostium bursae on eighth segment. Total length of pupa approxi- 
mately 6-7 mm. 

Adult (Fig. 14). As described (Stainton, 1861; Clarke, 1941). 


DISCUSSION 


The larvae of A. alstroemeriana should be confused with no other 
eastern North American Agonopterix species. Mature larvae can be 
distinguished from larvae of A. clemensella by the markings on the 
epicrania and the conspicuous pinacula, and from larvae of A. flavi- 
comella by the absence of a black cervical shield. The host plant also 
appears to be definitive—Conium maculatum is the only host record- 
ed for A. alstroemeriana, and A. alstroemeriana is the only oecopho- 
rid reported to feed on Conium. Larvae of A. clemensella, in fact, die 
when confined to foliage of C. maculatum (personal observation). 


42 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


7 8 


Fics. 4-8. Structures of larval head of A. alstroemeriana: 4, mandible, mesal view, 
showing condyle (C), lateral mandibular setae (LS), molar groove (MG), and first scis- 
sorial tooth (S); 5, lateral view of epicrania showing markings with ocelli (O); 6, 
hypopharyngeal complex, lateral view showing labial palps (LP) and spinneret (SP); 
7, spinneret, dorsal view, showing basal segment of labial palps (B) and silk pore (P); 
8, head, ventral view. 


VOLUME 37, NUMBER l 43 


13 AS 


Fics. 9-13. Structures of immature stages of A. alstroemeriana: 9, larval head, 
dorsal view, showing adfrontal setae (ADF1, ADF2), clypeal setae (C1, C2), frontal 
setae (F), frontal punctures (FP), first lateral setae (L1), first medial setae (M1), anterior 
setae (Al, A2, A3), lateral setae (L1) and posterior setal group (P1, P2, P3); 10, larval 
abdominal segments 1, 2, and 9, ventral view, showing lateral setae (L3), subventral 
setae (SV1, SV2, SV3) and ventral setae (V1); 11, pupal head and thorax, mesal view, 
showing antennae (A), labrum (L), mesothoracic legs (MOL), maxillary palps (MP) and 
maxillae (MX); 12, terminal abdominal segments of pupa showing male and female 
genital openings superimposed, ventral view, showing anal slit (An) and cremaster (C); 
13, fifth abdominal segments of pupa showing proleg scars and a portion of the fine 
setae covering most of the abdominal segments, ventral view. 


44 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Fic. 14. Adult of A. alstroemeriana. 


The adult is a typical Agonopterix in form and maculation, lacking 
the longitudinal streaks characterizing species of Depressaria. The 
only American oecophorid similar in coloration to A. alstroemeriana 
is Martyrhilda canella (Busck). Tufted labial palps and a simple distal 
process of the sacculus distinguish Agonopterix from Martyrhilda; 
moreover, M. canella has a dark head and lacks a brick red spot below 
the black patch on the forewings. The key to Agonopterix by Hodges 
(1974) may be amended as follows to include A. alstroemeriana: 


0. Head, thorax and basal streak of costa pure white, concolorous; discal cell of 
forewing with a black patch lying above a brick red spot __ A. alstroemeriana 
OP Notiasvabover o2.00 ogee Nee I 1 (start key) 


ACKNOWLEDGMENTS 


We thank J. G. Franclemont (Cormmell University, Ithaca, NY) for identifying A. al- 
stroemeriana, and E. R. Hoebeke (Cornell University) for comments on this manu- 
script. Alice Prickett (University of Illinois, Urbana-Champaign) patiently provided 
illustrative material. Specimens are on deposit in Cornell University Collection, Lot 
1023, sublot 41b. This work was supported by National Science Foundation grant DEB 
76-20114 to P. Feeny (Cornell University). 


VOLUME 37, NUMBER Il 45 


LITERATURE CITED 


BERENBAUM, M. 1981. Furanocoumarin distribution and insect herbivory in the Um- 
belliferae: plant chemistry and community structure. Ecology 62:1254—1266. 
BusH, A. 1908. A generic revision of American moths of the family Oecophoridae, 

with descriptions of new species. Proc. U.S. Natl. Mus. 35:187—207. 

CLARKE, J. F. G. 1941. Revision of the North American moths of the family Oeco- 
phoridae, with descriptions of new genera and species. Proc. U.S. Natl. Mus. 90: 
33-286. 

FORBES, W. T. M. 1923. The Lepidoptera of New York and neighboring states. Cor- 
nell University Agric. Exp. Station Mem. 68:1-479. 

GopFREY, G. L. 1972. A review and reclassification of the larva of the subfamily 
Hadeninae of America north of Mexico. U.S.D.A. Tech. Bull. 1450. 265 pp. 

HINTON, H.E. 1946. On the homology and nomenclature of the setae of lepidopterous 
larvae, with some notes on the phylogeny of the Lepidoptera. Trans. Roy. Entomol. 
Soc. London 97: 1-37. 

HopcEs, R. W. 1974. Gelechioidea, Oecophoridae. The Moths of America North of 
Mexico, Fascicle 6.2. E. W. Classey, Ltd., London. 142 pp. 

, F. M. Brown, D. R. Davis, D. C. FERGUSON, J. G. FRANCLEMONT, J. B. 
HEPPNER, L. D. MILLER, E. G. MUNROE & E. L. Topp. Checklist of the Lepi- 
doptera of America North of Mexico (in press). 

MacKay, M.R. 1972. Larval sketches of some microlepidoptera, chiefly North Amer- 
ican. Can. Entomol. Mem. 88:1-83. 

ScHUTzE, K. T. 1931. Die Biologie der Kleinschmetterlinge: Unter Besonderer 
Berucksichtigung Ihrer Nahrpflanzen und Erscheinungszeiten. Internat. Ent. Ver- 
ein E..V., Frankfurt. 235 pp. 

STAINTON, H. T. 1861. Depressaria pt. 1. Pt. vi in H. T. Stainton, P. C. Zeller & J. 
W. Douglas (eds.). Natural History of the Tineina. J. van Voorst, London. 

TOLL, S. 1964. Motyle: Lepidoptera. Oecophoridae. Pol. Zwiazek Klucze Oznacziana 
Owadow Pol. 27:1-174. 


Journal of the Lepidopterists’ Society 
37(1), 1983, 46-69 


GEOGRAPHIC DISTRIBUTION AND CHECKLIST OF THE 
BUTTERFLIES OF KERN COUNTY, CALIFORNIA 


KEN DAVENPORT 
6601 Eucalyptus Dr., #325, Bakersfield, California 93306 


ABSTRACT. A checklist of the 126 species of butterflies which are known to occur 
in Kern County, California is presented. Also considered is information concerning the 
distribution and flight periods for each species found regularly within the county 
boundaries. Briefly discussed are the natural features and geography of the area, con- 
sidering some of its characteristic plant life and climatic conditions. 


Butterflies and Kern County Geography 


Kern County is located in south-central California and embraces 
8064 square miles of very diversified territory, which includes the 
southern San Joaquin Valley, several mountain ranges and an arid 
portion of the Mojave Desert. Elevations range from 91 m (300 ft) 
above sea level at Lost Hills on the Valley floor to 2692 m (8831 ft) 
on the summit of Mt. Pinos (actually located on the Kern-Ventura 
County line), and 2583 m (8475 ft) on Owens Peak in the southern 
Sierra Nevada. 

The diversity in elevation, habitats and plant life support an un- 
usually large and varied butterfly fauna for an area the size of Kern 
County. At present, 126 species and a number of additional subspe- 
cies of butterflies have been collected within the county boundaries. 
Most of these are “residents” or regular visitors which establish tran- 
sient populations most years. A few are accidental to the region and 
occur here only as rare strays or migrants from adjacent areas where 
they are better established. These special cases will be discussed in 
the checklist portion of this paper. 

Kern County can be divided into three general areas geographical- 
ly. Each area supports its own plant and animal life, including the 
butterflies. 

(1) San Joaquin Valley: An arid lowland valley which is heavily 
used for agricultural purposes. Citrus and alfalfa (Medicago sativa L.) 
are among the predominate crops grown in the region, which are 
important to numerous kinds of butterflies. Rainfall is about 0.127 m 
(5 inches) per year, which is not enough to support much natural plant 
growth. What plants do grow here are adapted to low annual rainfall 
and very long, hot and dry summers. 

Most butterflies occurring in this region are species which are com- 
mon and widespread in the western United States. These butterflies 
readily adapt to man’s presence and influence and will be found in 
cities, residential areas, gardens, parks, and in agricultural fields. A 


VOLUME 37, NUMBER 1 AZ 


few other butterflies favor riparian habitats along the Kern River or 
Poso Creek. The Kern River drains the southern Sierra, including Mt. 
Whitney, and flows throughout the year (until it reaches Bakersfield 
and is diverted into irrigation canals and urban use) providing water 
for agricultural and urban development. Bakersfield with an unofficial 
population of some 250,000 is the largest city in the county and a 
number of smaller cities and towns lie nearby. 

Some of the species occurring on the Valley floor are butterflies 
which more normally would be expected out on arid stretches of the 
Mojave Desert. Among these are Pholisora libya, Pontia beckerii, 
Anthocharis cethura morrisoni and Danaus gilippus strigosus. These 
fly in undisturbed areas away from cities and agricultural fields and 
can be found in Atriplex wastelands, on alkali flats, in ravines or in 
swampy areas still found in otherwise dry arid country. Others like 
Pyrgus scriptura occur along irrigation ditches or roadsides. 

(2) Western Mojave Desert: The portion which lies in Kern 
County receives less than 0.254 m (10 inches) of rainfall in most years. 
As is true throughout the county, precipitation is strongly seasonal 
with most rainfall occurring in the winter and spring months. 

Vegetation is generally very sparse. What few plants grow in this 
area are adapted to highly arid conditions, high summer temperatures 
and sandy soils. Some common plants of the region are creosote bush 
(Larrea divaricata. Cav.), several varieties of rabbitbrush (Chryso- 
thamnus spp.), saltbrush (Atriplex spp.), buckwheats (Eriogonum spp.), 
Joshua tree (Yucca brevifolia Engelm. in Wats.) and several varieties 
of cacti. Various kinds of wildflowers can be found during the spring 
months. 

Most of the Kern County desert is flat and unremarkable, and few 
species of butterflies can be expected. Much better conditions exist 
where the desert and mountain regions meet. A great many desert 
butterflies occur on the arid east slope of the Sierra Nevada, in adja- 
cent canyons or in the high desert valleys. Walker Pass, elevation 1600 
m (5250 ft), in the southern Sierra is a particularly well known locality 
often visited by lepidopterists. Other lucrative areas for collectors are 
Red Rock Canyon, Jawbone Canyon and Kelso Valley. At least 70 
species of butterflies have been recorded from the Kelso Valley region 
alone, which makes this area exceptionally interesting. Several species 
more commonly encountered in the mountains can be found com- 
monly in desert washes out of their usual habitat. Most “desert” species 
of the region fly during the spring months of April and May, but a 
few species can be found through the summer and fall months as well. 

Two ‘rare’ desert species with very sporadic distributions in south- 
ern or southeastern California are Pseudocopaeodes eunus and Ple- 


48 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


bulina emigdionis. Both of these butterflies are locally common around 
Weldon, located along the south fork of the Kern River near Lake 
Isabella and on the northern edge of Kelso Valley. 

Still, the Kern County desert lacks the rich desert fauna of other 
deserts located to the south and east. Several “desert” species com- 
mon in Arizona or in southeastern California are encountered in this 
region only as rare strays or as small transient populations in years of 
favorable rainfall. 

(3) Montane areas: Mountains ring the San Joaquin Valley on three 
sides. These include the arid Temblor Range (part of the Coast Range) 
to the west; the Transverse Ranges (including Frazier Park and Mt. 
Pinos) to the south; the Tehachapi Mountains to the southeast; and 
the southern Sierra Nevada, Greenhorn and Piute mountain ranges to 
the east and southward. The Greenhorn and Piute ranges are actually 
subranges of the Sierra, but are best considered separately, because 
each of these three areas differ considerably ecologically and in but- 
terfly fauna. Most plants and animals (including the butterflies) of 
these mountain ranges are characteristic of the Upper Sonoran or 
Transition Life Zones. 

The mountains of Kern County are very rich in butterflies. For ex- 
ample, over 100 butterfly species have been collected within a radius 
of 15 miles from the town of Lake Isabella, which is adjacent to the 
southern Sierra, Greenhorn and Piute mountain ranges. Butterflies 
are found in these regions from late February or early March well 
into October or even November in some years. 

In the mountains grow several different species of conifers, includ- 
ing incense cedar (Calocedrus decurrens (Torr.)), ponderosa pine (Pi- 
nus ponderosa Lawson), Jeffrey pine (Pinus jeffreyi Grev. & Balf., in 
A. Murr.), sugar pine (Pinus lambertiana Doug]l.), some lodgepole 
pine (Pinus contorta murrayana Grev. & Balf.), digger pine (Pinus 
sabiniana Dougl.), white fir (Abies concolor (Gord. & Glend.)), red 
fir (Abies magnifica Murr.) and California juniper (Juniperus califor- 
nica Carr.). Along streams or in the mountain canyons are found white 
alder (Alnus rhombifolia Nutt.), sycamore (Platanus racemosa Nutt.), 
cottonwoods (Populus spp.) and several kinds of willows (Salix spp.). 
A number of species of oaks (Quercus) and buckwheats (Eriogonum) 
grow on the drier slopes. Many varieties of annual wildflowers and 
grasses are also found in these several mountain ranges. 

The Transverse Ranges are represented in Kern County by the Te- 
jon Mountains and are actually part of the Coast Range. Mt. Pinos and 
Frazier Mountain (the summit of which is actually in adjacent Ventura 
County) are both well over 2440 m (8000+ ft) elevation and are rich 
in butterflies. A number of species with very limited distribution (i.e., 


VOLUME 37, NUMBER Il 49 


Colias harfordii, Chalceria heteronea clara, Icaricia neurona and 
Speyeria coronis hennei) occur in this region. 

The Tehachapi Mountains rise to an elevation of 2435 m (7988 ft) 
on the summit of Double Mountain above Tehachapi Mountain Park 
near the town of Tehachapi. A trail leads to this mountain summit 
where is found the rare Speyeria egleis tehachapina and the some- 
what more common Speyeria coronis hennei. Only the latter species 
sometimes descends the lower slopes to the Mountain Park below. 
The highly sought Speyeria adiaste atossa once flew with Speyeria 
egleis tehachapina on the mountain summits but could be found de- 
scending to much lower elevations in shaded woodlands and along 
small streams. 

The Temblor Range is a low arid range located on the west side of 
the San Joaquin Valley. There are relatively few species in these 
foothills but one prized butterfly which does occur here is Mitoura 
siva mansfieldi, a subspecies distinguished by its dark green color- 
ation underneath. Despite an abundance of junipers in these foothills, 
mansfieldi is scarce and hard to find. 

The southern Sierra Nevada become very arid south of the Tulare 
County line. The range (not including the Greenhorns or Piutes be- 
cause these are being considered separately) continues south to near 
the edge of the Mojave Desert at Butterbredt Peak (also known as 
Butterbread Peak), which has an elevation of 1829 m (6000 ft). On 
these seemingly arid hillsides and hilltops fly the rare Pholisora al- 
pheus oricus and the southern Sierra subspecies of Papilio indra. 

The Kern County Sierra is too arid and low in elevation to support 
the array of boreal species found further north in the Canadian, Hud- 
sonian and Arctic-Alpine Life Zones of the truly high Sierra. How- 
ever, several of these high elevation species do occur just 10 to 15 
miles north of the county line and may occasionally stray southward 
to Kern County. Some of these may yet be found on Owens Peak or 
in adjacent mountain areas accessible only by well marked foot-trails. 
More study of this region is definitely needed. 

The Greenhorn Mountains receive heavy winter snows and are the 
most heavily wooded mountains in the county. The forested areas are 
poor butterfly habitats, but favorable places for lepidoptera exist in 
moist meadows, along roads or in various kinds of disturbed places. 
For many years this has been the well known habitat of Speyeria 
hydaspe viridicornis and the only known southern California locality 
for Clossiana epithore sierra, as cited in Emmel & Emmel (1973). 

The Piute Mountains are located south of the previous mountain 
range and Lake Isabella and are generally drier than the Greenhorns, 
yet somewhat surprisingly, are much richer for butterflies in general. 


50 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


KINGS COUNTY TULARE COUNTY INYO COUNTY 


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VENTURA COUNTY LOS ANGELES CoUNTY 


Fic. 1. Map of Kern County, California, showing geographical features and col- 
lecting localities. 


The highest summits are Piute Mountain at 2538 m (8326 ft) and Piute 
Peak at an elevation of 2570 m (8432 ft). Lepidopterists visit here 
rather regularly for Papilio indra, Mitoura siva juniperaria, Icaricia 
neurona, Speyeria egleis tehachapina, the very variable Thessalia 
leanira population of this region, Occidryas editha editha and a num- 
ber of other interesting species. 

Our knowledge of the Kern County fauna is incomplete, and ques- 
tions still remain to be answered. But, it would appear unlikely that 
many new species remain to be discovered in this area. A few will 
likely be found in the southern Sierra along the Tulare-Kern County 
line or in the Greenhorn Mountains. More work remains to be done 
in compiling distributional data, even on the floor of the San Joaquin 
Valley where unusual or unexpected species have turned up in recent 
years. 


Sources of Information for the Kern County Checklist 


I have actively collected in Kern County for 18 of the past 23 years, 
while Jim Brock has collected and studied the fauna of the region 
since 1968. In 1975 the two of us met and undertook the joint project 
of compiling a county list and defining the distribution of each species. 
Jim Brock was especially active in eastern Kern County, and his field 
work provided the basis for much of my own studies in recent years. 
Today, each of us has extensive collections of Kern County butterflies, 
which provide much of the basis and documentation for this paper. 


VOLUME 37, NUMBER lI ol 


During the course of our project an effort has been made to visit 
poorly collected localities overlooked by others. These include nu- 
merous places on the floor of the San Joaquin Valley, the Kern River 
Valley, Kelso Valley and in the southern Sierra. The result has been 
a wealth of new information and data to add to that already obtained 
by other workers who had already contributed heavily to this project. 

Emmel & Emmel (op. cit.) treated many of the species found in 
Kern County in their book on southern California butterflies and a 
number of scientific papers (too numerous to list) have dealt with 
some aspect of the region’s butterfly fauna. 

A number of lepidopterists have contributed data or information 
used in this paper. These include John Burns, Julian P. Donahue, 
John F. Emmel, Gary File, Rick Hewett, Weldon Kirk, Robert Lang- 
ston, John Luttrell, William W. McGuire, Ed Moran, James Mori, Paul 
Opler, Jerry Powell, Allen Rubbert, Ed Sampson and Charles Sek- 
erman. Past workers who contributed to our present knowledge in- 
clude John Adams Comstock, Charles M. Dammers and H. Morrison. 


Introduction to the Kern County Checklist 


The annotated checklist section of this paper considers 126 species 
for which there are definite records of capture from within the county 
boundaries. About 10 of these actually represent strays of species not 
normally found in the region or butterflies of questionable status which 
may not be a true part of the Kern County fauna. 

Following the main checklist is another listing of “doubtful or ques- 
tionable records,’ which considers a few additional species having 
been accredited to Kern County but which are based on questionable 
data, determinations or evidence. 

In listing the butterflies in the checklist, I have generally followed 
the nomenclature and phylogenetic order used in “A Catalogue/ 
Checklist of the Rhopalocera of America North of Mexico” (Lee D. 
Miller & F. Martin Brown, 1981, Mem. Lepid. Soc. No. 2). 

For species found in Kern County on a regular basis, information 
is presented regarding distribution and normal flight periods. Specific 
records are cited to add authority to the text and document previously 
unpublished information but are not given in the case of common 
species unless in some way significant. The dates given for flight 
periods are representative of when the butterfly normally occurs (or 
has been collected) and are not the result of an exhaustive study of 
the collection data of the various contributors. While earlier and later 
collection data undoubtedly exist for many of the butterflies consid- 
ered, the dates given provide a fairly accurate picture of the year-to- 


52 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


year flights for most species or subspecies. Of course, more data are 
needed on several of the less commonly encountered butterflies. 

Also considered in the text are unusual populations, comments re- 
garding taxonomy, or other information of interest not published or 
generally well known. An effort has been made not to simply dupli- 
cate or restate information already published somewhere else in the 
literature. 

The varied topography of Kern County supports a number of un- 
described or atypical populations. Some of these will undoubtedly be 
described as new subspecies by other lepidopterists in the near fu- 
ture. Any mention or description of these given in this paper is merely 
intended to show how a specific population differs from other de- 
scribed known populations. These comments are not intended as a 
formal description of any “new” subspecies. Which of these unusual 
or atypical populations warrant names or recognition in the nomen- 
clature will be left to the taxonomists and specialists in their respec- 
tive fields. 

To conserve space in the checklist a distribution code is used to 
represent various general localities where the butterflies can be found. 
Such a code represents the names of the major valleys, canyons and 
mountain ranges. The names of cities and towns or the citations of 
specific locality data will not be coded. The code is as follows: 


(1) Code for the major valleys or lowlands: 
SJV: The San Joaquin Valley, often including Bakersfield. 
KRV: The Kern River Valley, often including Kernville, Lake Isabella and Wel- 
don. 
Code for the desert regions: 
MD: The Mojave Desert. 
RRC: Red Rock Canyon and vicinity. 
WP: Walker Pass, often on the dry eastern slope of the Sierra. 
JC: Jawbone Canyon, located at the south end of the Sierra. 
KV: Kelso Valley, often including Sageland and arid foothills. 
(3) Code for the mountain regions: 
MTNS: Mountains, distribution general throughout most of the mountain areas 
of Kern County. 
SNM: Sierra Nevada Mountains, including the area along the Kern-Tulare County 
line southward to Butterbredt Peak. 
GM: Greenhorn Mountains, including Cedar Creek and Mountain Park. 
PM: Piute Mountains, including areas adjacent to Bodfish and Havilah; also 
includes Hooper Hill. 
TM: Tehachapi Mountains, includes Tehachapi Mountain Park and the Cal- 
iente region. 
TJM: Tejon Mountains, including Frazier Park and Mt. Pinos. 
TR: Temblor Range, including McKittrick and Taft. . 
KRC: Kern River Canyon, including Miracle and Democrat Hot Springs and the 
area around Richbar. 


(2 


—V~ 


Specific records given in the text are those of the author unless 
otherwise stated. 


VOLUME 37, NUMBER 1 53 


AN ANNOTATED CHECKLIST OF THE BUTTERFLIES OF 
KERN COUNTY, CALIFORNIA 


HESPERIIDAE 
1. Polygonus leo leo (Gmelin). 


Jim Brock collected a single worn specimen of this skipper 5 miles east of Caliente 
on 6 IX 73. This record probably represents a wind blown stray. It is almost certainly 
not a regular member of the county fauna. 


2. Thorybes pylades (Scudder). 

Distribution: SNM, GM, TM, TJM. Flight: 13 VI to 23 VI. 

Widely distributed but rarely encountered. Most records are from the Frazier Park 
region where pylades probably first appears in May. 


3. Erynnis brizo lacustra (Wright). 
Distribution: GM, PM, TM, TJM. Flight: 3 IV to 30 VI. 


4. Erynnis propertius (Scudder & Burgess). 
Distribution: MTNS. Flight: 11 III to 11 VII. 


5. Erynnis tristis tristis (Boisduval). 

Distribution: KRV, SJV, PM (Hooper Hill). Flight: 25 IV to 24 IX. 

Common at Bakersfield in 1962 and 1963, tristis has since become exceedingly scarce 
in the southern San Joaquin Valley. 


6. Erynnis pacuvius (Lintner). 


John Burns examined material of this species from Kern County (in 1980) and de- 
termined that two subspecies occur here: 


a. Erynnis pacuvius lilius (Dyar). 
Distribution: GM, PM. Flight: 11 VI to 21 VII. 


b. Erynnis pacuvius callidus (Grinnell). 


Distribution: TM, TJM. Flight: 31 V to 21 VII. 
Generally callidus is found further south and southwest than lilius. Sympatry has 
not been observed. 


7. Erynnis funeralis (Scudder & Burgess). 


Distribution: SJV, KRV, PM, MD. Flight: 10 IV to 9 X. 
Common throughout most of southern California, funeralis is rare in Kern County. 
Like E. tristis, this species was common in Bakersfield in 1962 and 1963. 


8. Erynnis persius persius (Scudder). 


The status of this widely distributed Erynnis in Kern County is presently unknown. 
It is included on the county list on the basis of a single specimen collected by Jerry 
Powell at Delano on 12 VII 55 (Burns, 1964). John Burns (pers. comm.) suspects that 
persius may be “resident” within the county boundaries and is probably much more 
widely distributed here than the lone record would suggest. It has been collected at 
Big Meadow in the southern Sierra, Tulare County, which is just a few miles north of 
the county line. 


9. Pyrgus scriptura (Boisduval). 

Distribution: Western SJV. Flight: 27 II to 3 X. 

Most of our records are from Buttonwillow and Lost Hills. Scriptura appears to be 
absent from agricultural areas around Bakersfield and the eastern side of the San Joa- 
quin Valley. Very local. 


54 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


10. Pyrgus communis albescens Plotz. 
Distribution: SJV, KRV, KV, MD, MTNS, at lower elevations. Flight: 25 II to 9 XI. 


11. Heliopetes ericetorum (Boisduval). 
Distribution: SJV, KRV, KV, JC, MTNS. Flight: 30 IV to 9 X. 


12. Pholisora catullus (Fabricius). 


Distribution: SJV, KV, PM, east slope of Breckenridge Mountain. Flight: Mountains 
and desert: 3 IV to 7 VI. SJV: 20 VIII to 5 IX. Probably double brooded in the southern 
San Joaquin Valley. The mountain and desert populations appear to be single brooded. 

Catullus appeared to be absent from the floor of the southern San Joaquin Valley 
until a small population was found in a swampy area adjacent to the Kern River flood 
canal on the Tule Elk State Reserve on 20 VIII 81. 


13. Pholisora libya (Scudder). 


Two variable and geographically separated populations occur in Kern County. Out 
on the Mojave Desert (including Homestead and Jawbone Canyon) and on the east 
slope of the Sierra Nevada there is a rather mixed population which shows character- 
istics of nominate libya and the larger lena (Edwards) but which tends toward lena 
(John F. Emmel, pers. comm.). Individuals of this eastern Kern County population are 
frequently encountered which have the undersurface of the hind wings entirely colored 
with white scales. 

In the western San Joaquin Valley (Maricopa, Taft, Buttonwillow, McKittrick and 
Lost Hills) is an undescribed subspecies (Emmel & Emmel, 1973) distinguished by its 
larger size and washed-out appearance of the underside of the hind wings. 

Flight of P. libya/lena: 6 V to 26 V; 21 VIII to 30 IX. 

Flight of SJV subspecies: 28 IV to 18 V; 20 VI to 12 IX. The spring brood of the SJV 
populations is the heaviest flight; later broods are unreliable and made up of few 
individuals. This is in contrast with the desert population which has its heaviest flight 
in the late summer and early fall. 


14. Pholisora alpheus oricus Edwards. 


Distribution: SNM (Butterbredt Peak). One record for KV. Flight: 21 V to 15 VI. 
Probably appears in late April. 

A rare butterfly throughout most of its range, oricus is locally abundant on the eastern 
and southern slopes and canyons of Butterbredt Peak where the Atriplex foodplant 
grows. 


15. Copaeodes aurantiaca (Hewitson). 


Distribution: Homestead, WP, KV, Caliente Canyon. Flight: 1 V to 22 X. 

Aurantiaca breeds in small numbers in the desert areas and in low arid mountain 
canyons. Tom Rubbert collected a specimen in the San Joaquin Valley at Bakersfield 
on 17 VI 50. Allen Rubbert has a specimen from the same locality which lacks data as 
to the date of capture. 


16. Hylephila phyleus (Drury). 
Distribution: MD, SJV, KRV, Caliente region. Flight: 8 IV to 12 XII. 


17. Pseudocopaeodes eunus eunus (Edwards). 


Distribution: Weldon, Onyx and probably elsewhere in KRV. Formerly known from 
S]V where relict populations may still exist. Flight: 14 VI to 21 VIII in KRV. 

John Adams Comstock (Comstock, 1927) reported that eunus was known from the 
southern San Joaquin Valley at that time. C. M. Dammers collected at least one female 
specimen (now in the Los Angeles Museum) at Panama (near Bakersfield) on 18 VIII 
29. This population may have disappeared with the destruction of its saltgrass habitat 
due to urban and agricultural development of the region. At least no recent records for 


VOLUME 37, NUMBER l 55 


this species in the San Joaquin Valley are known to the author. However, eunus can 
still be found commonly in saltgrass habitats at Weldon and Onyx where it appears to 
fly continuously throughout the summer months. Ron Leuschner discovered this pop- 
ulation on the Kern River Preserve at Weldon on 13 VI 81. 


18. Hesperia juba (Scudder). 

Distribution: SNM, GM, PM, TM, TJM, WP, KV. Usually uncommon. Flight: 9 V to 
20 VI; 11 IX to 14 X. More common in the fall. 
19. Hesperia comma (Linnaeus). 


Two distinct populations occur in Kern County: 


a. Hesperia comma harpalus (Edwards). 
Distribution: KRV, KV, SNM, WP, PM (east slope). Flight: 19 VI to 5 IX. 


b. Hesperia comma near tildeni Freeman. 


Distribution: PM (west slope), KRC, TM, GM, TJM. Flight: 1 VII to 9 X. 

This population of comma is smaller and lighter than harpalus. How this popu- 
lation is actually related to other Sierran and Coast Range populations is presently 
under study by William W. McGuire. The Sierran subspecies yosemite Leussler 
probably does not occur in Kern County. 


20. Hesperia columbia (Scudder). 


Distribution: Kernville, PM, TJM. Flight: 31 III to 24 V; 12 IX. Ray Stanford col- 
lected a specimen on the summit of Mt. Pinos on 29 VI 63, an extreme collection date. 
Most records are for May. 

Columbia is intensely local and uncommon and few records exist for Kern County. 
It has been collected in numbers at two localities in Ventura County adjacent to the 
Kern County line. It should be found at several additional localities as more favorable 
habitat areas are explored in the future. 


21. Hesperia lindseyi Holland. 


Distribution: GM, PM, TM, TJM. Flight: 14 V to 12 VII. 
Usually uncommon and local in occurrence. 


22. Polites sabuleti sabuleti (Boisduval). 

Distribution: SJV, KRV, KV, Havilah, TM, TJM. Flight: 4 V to 1 XI. Probably first 
appears in April. 

Individuals which resemble chusca (Edwards) are regularly found in most county 
populations. 
23. Polites sonora sonora (Scudder). 

Distribution: GM, TM. Flight: 13 VI to 11 VII. 


24. Atalopedes campestris campestris (Boisduval). 
Distribution: MD, SJV, KRV, KRC, PM, TM, TJM. Flight: 28 III to 13 XI. 


25. Ochlodes sylvanoides sylvanoides (Boisduval). 
Distribution: MTNS. Flight: I VII to 9 X. 


26. Ochlodes agricola (Boisduval). 
Three subspecies occur in Kern County: 


a. Ochlodes agricola agricola (Boisduval). 
Distribution: TM, TJM. Flight: Late May to early July. 


56 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


b. Ochlodes agricola verus (Edwards). 
Distribution: Havilah, PM, KV. Flight: 26 V to 21 VII. 


ce. Ochlodes agricola nemorum (Boisduval). 


Distribution: Kernville, SNM, GM. Flight: 19 V to 7 VII. 
Nemorum blends with verus on the northern edge of the Piute Mountains around 
Bodfish. 


27. Paratrytone melane melane (Edwards). 

Distribution: KRC along Clear Creek at Miracle Hot Springs. Flight: 14 V to 21 V; 
10 VIII. 

Widespread elsewhere in southern California, this skipper thus far has been found 
only at the above mentioned locality where it is uncommon. 


28. Lerodea eufala (Edwards). 


Distribution: MD, SJV, KRV. Flight: 23 III to 12 XII. Most records are from August 
through October. 


MEGATHYMIDAE 
29. Megathymus coloradensis martini Stallings & Turner. 


Distribution: SNM (arid slopes), WP, KV. Flight: 3 IV to 25 V. 
This large skipper is rarely taken in numbers but probably has a wider range out on 
the desert floor where stands of the Yucca larval foodplant are rather common. 


PAPILIONIDAE 
30. Battus philenor (Linnaeus). 


This is not a regular member of the county fauna and apparently occurs only as a 
stray. There is only one definite record: 13 miles west of Shafter (a male on alfalfa); 
30 VI 57; collected by Paul Opler. Which subspecies this record represents is ques- 
tionable. Either the nominate subspecies or northern California hirsuta (Skinner) could 
stray into the county at times. 


31. Papilio rudkini Chermock & Chermock. 


Distribution: SNM (Butterbredt Peak), one record for RRC by Jim Brock (5 IV 74). 
Flight: 5 IV to 26 V. 

This desert swallowtail is an uncommon find in Kern County, but there are several 
records for adults and larvae (on Tauschia parishii (C. & R.)) on Butterbredt Peak. 
(Thamnosma montana Torr. & Frem. is the usual host in most of its range.) Many 
lepidopterists believe that rudkini is probably a subspecies of Papilio polyxenes Fa- 
bricius. 


32. Papilio zelicaon zelicaon Lucas. 
Distribution: SJV, MTNS. Flight: 25 II to 25 X. 


33. Papilio indra phyllisae J. Emmel. 


Distribution: SNM (Butterbredt Peak and vicinity), PM (Piute Mountain Vista; also 
1 mile west of Kelso Valley Road Summit at east end of Piute Mountains). Flight: 13 
IV to 30 VII. This subspecies has a small second brood which flies during the month 
of July. 

Though presently known from but three localities in the county, phyllisae will un- 
doubtedly be found on several of the higher peaks in the southern Sierra Nevada where 
the Tauschia parishii foodplant is widely distributed. 


34. Pterourus rutulus rutulus (Lucas). 
Distribution: SJV, KRV, MTNS. Flight: 8 III to 25 X. 


VOLUME 37, NUMBER l 57 


35. Pterourus multicaudata (Kirby). 


Distribution: GM (Cedar Creek), PM, TM, TJM. Flight: 29 V to 30 VII. Probably 
appears in early May on east slope of Breckenridge Mountain near Havilah. 

Multicaudata tends to be local in occurrence and uncommon but is sometimes locally 
abundant. It prefers canyon bottoms with small streams where oak woodland and co- 
niferous forest meet. , 


36. Pterourus eurymedon (Lucas). 
Distribution: MTNS. Flight: 24 IV to 21 VII. 


PIERIDAE 
37. Neophasia menapia menapia (Felder & Felder). 
Distribution: GM, PM, TM. Flight: 9 VII to 20 VIII. 


38. Pontia beckerii (Edwards). 


Distribution: SJ)V, KRV, MTNS, MD. Flight: 22 III to 3 XI. 

One of the surprises of this survey was finding a large population of this butterfly on 
the floor of the San Joaquin Valley. A few beckerii adults had been collected around 
Bakersfield prior to 1981, but it was assumed that these individuals represented strays 
from other areas. Actually, beckerii appears to be well established in the foothills 
northeast of Bakersfield (southeast edge of Hart Park, 20 VI 72; 6 to 16 VI 81) and north 
of Oildale (Poso Creek 8 miles north of Oildale, 18 V to 26 VI 81). It was also abundant 
along the Glennville—Woody Road 8 to 20 miles north of Oildale on 26 VI 81. The 
bladderpod plant, Isomeris arborea Nutt., grows commonly in this portion of the San 
Joaquin Valley and is likely used as a larval host. 


39. Pontia sisymbrii sisymbrii (Boisduval). 
Distribution: MTNS, WP, KV, Kernville. Flight: 25 II to 30 V. 


40. Pontia protodice (Boisduval & LeConte). 
Distribution: Entire county. Flight: 12 II to 24 XI. 


41. Artogeia rapae (Linnaeus). 


Distribution: Entire county except arid portions of desert. Flight: All months, but 
mostly from March through October. 


42. Euchloe hyantis (Edwards). 


Two sets of populations occur in Kern County: 


a. Euchloe hyantis lotta (Beutenmuller). 


Distribution: MD, SNM, PM (east slope), TM (arid southern and eastern slopes). 
Flight: 12 III to 15 V. 


b. Euchloe hyantis (Edwards) ssp. “Mt. Pinos block segregate.” (Opler, 1968, 1969; 
also Emmel & Emmel, 1973) 


Distribution: TJM. Flight: 5 V to 8 VI. 
This population is uncommon and rarely collected in numbers. 


43. Anthocharis cethura morrisoni Edwards. 


Distribution: Western SJV, TR, SNM, PM (Hooper Hill and arid eastern slope), KRC 
(Miracle Hot Springs), KV, WP, RRC. Flight: 19 II to 15 V. Certain populations (KRC 
and SJV) have been noted to forego emergence in years of unfavorable rainfall. 

This subspecies is distinguished by the dark green coloration of the hind wings. 
Most females of the lower Kern River and valley populations lack the “orange-tip” 
though females from eastern Kern County often do. 


58 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


44. Anthocharis sara sara Lucas. 


Distribution: KRV, TR, MTNS, KV. Flight: 17 II to 21 VI. The early spring brood 
(reakirtii Edwards) is the most common. The larger and lighter second brood is rela- 
tively scarce. 


45. Falcapica lanceolata lanceolata (Lucas). 


Distribution: SNM, GM, PM, TJM, KRC, WP, Erskine Creek near Lake Isabella. 
Flight: 28 II to 19 VI. 

Though generally considered to be a rare species, lanceolata is often abundant along 
Erskine Creek and in the Piute Mountains just south of Bodfish. 


46. Colias eurytheme Boisduval. 


Distribution: Entire county. Flight: 20 II to 5 XI. 
This species is often a serious economic pest in the San Joaquin Valley. 


47. Colias harfordii Hy. Edwards. 


Distribution: KV, PM, TM, TJM. Flight: 28 V to 26 VIII. Another flight in late 
September and October is probable since harfordii was observed and collected in 
numbers on the north slope of Frazier Mountain in Ventura County (just across the 
county line) on 9 X 81. 

Most records for the county are from Frazier Park and Mt. Pinos. Elsewhere, it tends 
to be very rare and unreliable. The northernmost records for harfordii are from the 
Piute Mountains (near Liebel Peak, 21 VII 78) and Kelso Valley (the area around 
Sageland) where the butterfly occurs in June. 


48. Zerene eurydice (Boisduval). 


Distribution: TM, TJM. Flight: 23 VI to 11 VII. 

Though common in much of southern California, this butterfly is scarce in Kern 
County. It is uncommon in the Transition Zone of Tehachapi Mountain Park and around 
Frazier Park. It can sometimes be fairly common on dry hillsides near the McGill 
Campground on Mt. Pinos. Eurydice appears to be single-brooded at these localities 
though multiple-brooded elsewhere in southern California. 


49. Zerene cesonia cesonia (Stoll). 


This species is uncommonly found in Kern County, but it evidently establishes tran- 
sient breeding populations out on the desert floor. Many were seen in Jawbone Canyon 
and on Tom’s Hill in April and May of 1978. I know of two definite records: Weldon 
Kirk collected a specimen at Tehachapi Mountain Park in August of 1962 (specific date 
unknown); and I have a record for Koehn Dry Lake, 18 IV 78. 


50. Phoebis sennae marcellina (Cramer). 


This is not a regular member of the county fauna, though it may occasionally breed 
on Cassia plants out in the Mojave Desert. Ed Sampson collected a single male near 
Arvin in the San Joaquin Valley on 10 III 68. It has been collected on Frazier Mountain 
just south of the county line in Ventura County. 


dl. Eurema mexicana (Boisduval). 


This is another species which reaches the area periodically and which may establish 
transient populations on Cassia plants out on the desert floor. There are definite records 
for Frazier Park, Bodfish, Weldon and Lakeview (in the San Joaquin Valley) where Ed 
Sampson collected a specimen on 16 V 66. Most of the other records are also for the 
month of May. 


52. Abaeis nicippe (Cramer). 


Distribution: MD (including Ridgecrest) with other records of strays or transients 
from north of Kernville, Bodfish, PM, WP, JC and Caliente Canyon. Flight: 6 III to 
9 X. 


VOLUME 37, NUMBER 1 59 


This species may not be permanently established in Kern County and may need to 
periodically reestablish itself here after cold winters. Records for early March in 1978 
indicate that it does successfully overwinter at times. 

53. Nathalis iole Boisduval. 


This is another species which enters the region only occasionally. Records exist for 
the Mojave Desert, adjacent mountain ranges (PM, TM, TJM) and even the San Joaquin 
Valley. Most of these records are for May or early June. 

LYCAENIDAE 
54. Tharsalea arota arota (Boisduval). 
Distribution: MTNS, KRV. Flight: 20 V to 10 VIII. 


55. Gaeides xanthoides xanthoides (Boisduval). 
Distribution: MTNS, KRV, KV. Flight: 21 V to 4 VIII. 


56. Gaeides gorgon (Boisduval). 
Distribution: MTNS, KV. Flight: 14 V to 19 VI. 


57. Chalceria heteronea clara (Hy. Edwards). 

Distribution: PM, TM, TJM. Flight: 22 VI to 26 VII. 

Clara has a very restricted range in southern California where it tends to be scarce 
and local in occurrence. Many former habitat areas have disappeared because of human 
influence. It is still common in certain canyons and washes in the Tejon Mountains. 
Most records are from around Lebec and Frazier Park. 

58. Epidemia helloides (Boisduval). 

Distribution: SJV (rare), KRV, Havilah, KRC, Paris-Loraine, Frazier Park. Flight: 30 
IV to 1 XI. 

59. Habrodais grunus grunus (Boisduval). 

Distribution: MTNS. Flight: 26 VI to 4 IX. 


60. Atlides halesus estesi Clench. 


Distribution: SJV, KRV, TM, TJM, TR. Flight: 10 III to 21 X. 

This spectacular butterfly is sometimes abundant in residential areas of Bakersfield, 
along the Kern River at Hart Park and in the Caliente region. Usually, it is rather 
scarce. 

61. Satyrium behrii behrii (Edwards). 


Distribution: KV, PM (east slope), TM, TJM. Flight: 14 VI to 10 VII. 


62. Satyrium californica (Edwards). 
Distribution: MTNS. Flight: 3 VI to 20 VII. 


63. Satyrium sylvinus (Boisduval). 

Sierran S. sylvinus sylvinus may be the subspecies found in the Greenhorn Moun- 
tains. Two other subspecies occur in Kern County: 

a. Satyrium sylvinus dryope (Edwards). 


Distribution: KRC, TM, TJM. Flight: 1 VI to 4 VIII. 
Dryope is usually found at lower elevations than the next subspecies. It is also 
less common than desertorum. 


b. Satyrium sylvinus desertorum (Grinnell). 


Distribution: KRV, MTNS. Flight: 29 V to 10 VIII. 
Sylvinus from Havilah and the Greenhorn Mountains appear darker underneath 


60 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


than desertorum from the Tehachapi Mountains. These need further study before 
they can be properly placed taxonomically. 

64. Satyrium auretorum spadix (Hy. Edwards). 
Distribution: TR, GM, TM, TJM. Flight: 8 VI to 30 VII. 


65. Satyrium tetra (Edwards). 
Distribution: PM, TM, TJM. Flight: 19 VI to 26 VII. 


66. Satyrium saepium saepium (Boisduval). 
Distribution: MTNS. Flight: 14 V to 26 VII. 


67. Callophrys dumetorum dumetorum (Boisduval). 

Distribution: SNM, PM, KRC, TJM, KV. Flight: 20 II to 22 V. 

Many Kern County populations seem atypical and warrant further study. 
68. Mitoura spinetorum (Hewitson). 


Distribution: MTNS. Flight: 30 IV to 17 IX. 

This hairstreak is rarely found in numbers. It is sometimes common on flowers or 
along streams at Tehachapi Mountain Park or in the Caliente region. Gary File has one 
record for Hart Park in the San Joaquin Valley, 17 IX 73. 

69. Mitoura nelsoni nelsoni (Boisduval). 


Distribution: GM. Flight: 6 VI to 17 VII. 
70. Mitoura siva (Edwards). 


Two subspecies occur in Kern County: 


a. Mitoura siva juniperaria Comstock. 
Distribution: SNM, Bodfish, PM, TJM. Flight: 3 IV to 30 VII. 


b. Mitoura siva mansfieldi Tilden. 
Distribution: TR. Flight: Late March to 17 IV. 


71. Incisalia augustus iroides (Boisduval). 
Distribution: MTNS. Flight: 19 III to 6 VI. 


72. Incisalia eryphon eryphon (Boisduval). 
Distribution: GM, PM. Flight: 11 VI to 9 VII. 


73. Strymon melinus pudica (Hy. Edwards). 
Distribution: Entire county. Flight: 3 III to 7 X. 


74. Brephidium exilis (Boisduval). 

Distribution: SJV, KRV, MD, MTNS (usually at low elevations). Flight: 27 II to 
15 XI. 
75. Leptotes marina (Reakirt). 

Distribution: SJV, KRV, MD, MTNS. Flight: 3 IV to 3 X. 


76. Hemiargus ceraunus gyas (Edwards). 


This butterfly appears to periodically establish itself in the lower mountain canyons 
of the Tehachapi and Piute Mountains. It has been collected at Miracle Hot Springs 
in Kern Canyon, south of Bodfish in the Piute Mountains, 5-10 miles east of Caliente, 
3 miles north of Twin Oaks and even in the San Joaquin Valley where a fresh female 
was collected along the Kern River at Hart Park, 23 IX 1982. Flight: 21 V to 23 IX. 


VOLUME 37, NUMBER 1 61 


77. Hemiargus isola alce (Edwards). 


I collected a single fresh female near Buttonwillow in the San Joaquin Valley on 2 
IX 80. Mesquite (Prosopis) still grows in a few places on the valley floor and out on 
the Mojave Desert and may support populations of this butterfly. 


78. Everes comyntas comyntas (Godart). 


This species may be a “resident” somewhere in the county but no populations are 
presently known. I found a small transient population along the Kern River at Hart 
Park (4 VIII 70) which was subsequently destroyed by flooding. 


79. Everes amyntula amyntula (Boisduval). 
Distribution: TM, Frazier Park. Flight: 18 III to 5 VII. 


80. Celastrina ladon echo (Edwards). 
Distribution: GM, PM, KRC, TM, TJM. Flight: 4 III to 14 VII. 


81. Euphilotes battoides (Behr). 


Two subspecies occur in Kern County: 


a. Euphilotes battoides bernardino (Barnes & McDunnough). 


Distribution: MTNS (including E] Paso Mtns. and TR), WP, KV, JC. Flight: 26 V 
to 26 VII. 

Eastern Kern County desert populations of battoides have been called martini 
(Mattoni) but are better viewed as atypical bernardino, according to John F. Emmel 
(pers. comm.). 


b. Euphilotes battoides comstocki (Shields). 


Distribution: Piute Mountain Vista, TM. Flight: 18 VII to 22 VII. Adults undoubt- 
edly fly into the month of August. 

Jim Brock recently discovered a large colony of comstocki on Piute Mountain Vista 
(or Lookout). Prior to his discovery this subspecies was known in California from 
only a few specimens collected in the Tehachapi Mountains (22 VII 18) by John A. 
Comstock. Adults are associated with a yellow flowered Eriogonum. 


82. Euphilotes enoptes (Boisduval). 

The taxonomic arrangement of this species in Kern County is very perplexing and 
open to considerable subjective opinion. Oakley Shields (1977) felt at that time the 
variable and atypical populations found in the southern Sierra, Piutes, Tehachapi and 
Mt. Pinos areas should best be viewed as forms of E. enoptes enoptes. Since Shields 
wrote his paper other populations have been discovered which do not appear to fit into 
this concept. My presentation of this species is tentative pending further study by other 
workers. 


a. Euphilotes enoptes enoptes (Boisduval). 
Distribution: MTNS. Flight: 24 IV to 26 VII. 


b. Euphilotes enoptes tildeni (Langston). 


Distribution: TR. Flight: Late August to early September. Jim Brock has one record 
hon 2? LIL 77: 

A large population exists on dry hillsides in the Temblor Range. Most records are 
from along Highway 58 near the Kern-San Luis Obispo County line. 


c. Euphilotes enoptes mojave (Watson & Comstock). 
Distribution: SNM (east and south slopes), MD, KV, JC. Flight: 9 IV to 26 V. 


62 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


d. Euphilotes enoptes (Boisduval) ssp. Butterbredt Peak population. 


Distribution: SNM (Butterbredt Peak and vicinity), PM (east slope), KV. Flight: 
Late April to early June. 

This population is similar to mojave (which practically “surrounds” the Butter- 
bredt Peak locality) but is much larger and is associated with Eriogonum nudum 
Dougl. ex Benth. which is used as the larval host (John F. Emmel, pers. comm.). 


e. Euphilotes enoptes (Boisduval) ssp. Fall-flying population. 


Distribution: Some of the same areas where enoptes enoptes flies in the spring. 
Hooper Hill, PM, KV, SNM, JC. Flight: 31 VIII to 9 X. 

This enoptes is distinguished by bolder black spots on the ventral side and by 
darker suffusion of the forewings beneath. John Emmel reports that this same “sub- 
species” occurs on the north slopes of other southern California mountain ranges 
and in Inyo County. 


83. Euphilotes pallescens elvirae (Mattoni). 


Distribution: SNM (arid slopes), WP, PM (Harris Grade), Tehachapi. Flight: Mid- 
June to late September. 

Shields (1977) notes that many specimens from Walker Pass resemble pallescens 
(Tilden & Downey). 


84. Philotiella speciosa speciosa (Hy. Edwards). 


Distribution: RRC, Randsburg and other locations in MD. No populations are pres- 
ently known at Havilah (the type locality) or in the southern San Joaquin Valley of 
Kern County. Flight: Mid-April to 13 V. 


85. Glaucopsyche piasus piasus (Boisduval). 


Distribution: SNM, GM, PM, TJM, KV. Flight: 3 IV to 18 VI. 
A huge population exists near Sageland in Kelso Valley where adults fly in desert 
washes (in association with a Lupinus spp.) out of the usual montane habitat. 


86. Glaucopsyche lygdamus (Doubleday). 


Three distinctive populations occur in Kern County: 


a. Glaucopsyche lygdamus australis Grinnell. 
Distribution: TM, TJM. Flight: 20 V to 14 VII. 


b. Glaucopsyche lygdamus columbia (Skinner). 

Distribution: SNM, GM. Flight: March to early May. 

Langston (1969) has columbia extending the length of the Sierra Nevada; whereas, 
incognitus Tilden is restricted to the central Coast Ranges. Kern County columbia 
has larger black spots underneath than the more northerly populations of this sub- 
species. 


c. Glaucopsyche lygdamus (Doubleday) ssp. 


Distribution: KRC, PM, WP, KV. Flight: 12 III to 26 V. 

This population is distinguished by bold black spots on both wings underneath 
and a smaller size than columbia. The females often have considerable blue scaling 
above and resemble australis in this respect. This “subspecies” appears to be in- 
termediate between the two previously discussed subspecies. 


87. Lycaeides melissa paradoxa (Chermock). 
Distribution: TJM, TM, KRV, KV. Flight: 3 IV to 3 X. 


88. Plebejus saepiolus saepiolus (Boisduval). 
Distribution: SNM (Fay Creek), GM, PM. Flight: 19 VI to 11 VII. 


VOLUME 37, NUMBER Il 63 


Some Kern County material resembles southern California hilda (Grinnell & Grin- 
nell). 


89. Plebulina emigdionis (Grinnell). 


Distribution: TJM including San Emigdio Canyon (the type locality), KRV including 
Weldon. Flight: 23 IV to 11 IX. 
Emigdionis is very abundant on Atriplex at Weldon (Paul’s Place). 


90. Icaricia icarioides evius (Boisduval). 
Distribution: MTNS, KV. Flight: 3 IV to 14 VII. 


91. Icaricia acmon acmon (Westwood & Hewitson). 
Distribution: Entire county. Flight: 21 III to 9 X. 


92. Icaricia lupini (Boisduval). 


Two subspecies occur in Kern County: 


a. Icaricia lupini monticola (Clemence). 
Distribution: MTNS, WP, KV, KRV. Flight: 10 IV to 14 VII. 


b. Icaricia lupini chlorina (Skinner). 

Distribution: TM, TJM. Flight: May to early July. 

Populations of chlorina are very uncommon. Paul Opler found it in the hills west 
of Lebec (9 VI 57) associated with Eriogonum nudum. 


93. Icaricia neurona (Skinner). 


Distribution: SNM (Pacific Crest Trail 6-9 miles north of Weldon), KV (wash 1 mile 
south of Sageland), Erskine Creek nr. Lake Isabella, PM (1-2 miles south of Bodfish 
and on Hooper Hill), TM, TJM. Flight: 30 IV to 26 VIII. Possibly flies well into 
September in some years. In some areas two or more broods are indicated. 

This blue is often locally abundant in the Piute Mountains just south of Bodfish 
where it frequents canyon bottoms and roadsides. 


RIODINIDAE 
94. Apodemia mormo (Felder & Felder). 


Populations of this species vary considerably, and much confusion exists about the 
taxonomy of the species. Kern County populations are perplexing and await further 
study by a specialist. At least three distinct groups exist: 


a. Apodemia mormo nr. mormo (Felder & Felder). 


Distribution: E] Paso Mountains, Homestead, Randsburg, JC. Flight: 18 IV to 13 
VWeLZIUVITT to 9 X. 

Fall flying specimens from Homestead and Jawbone Canyon closely resemble 
deserti Barnes & McDunnough. These populations fly in association with Eriogonum 
inflatum (Benth.) S. Stokes or with Eriogonum heermannii Dur. & Hilg. (Jim Brock, 
pers. comm.). 


b. Apodemia mormo nr. tuolumnensis Opler & Powell. 


Distribution: TR, SNM, KV, PM, TJM. Flight: 30 VII to 3 X. 
Is associated with Eriogonum wrightii Torr. Locally common. 


c. Apodemia mormo virgulti (Behr). 


Distribution: SNM, KV, PM, WP. Flight: 13 IV to 26 V. Dark form: 27 VIII to 2 X. 

Individuals associated with Eriogonum fasciculatum Benth. (Jim Brock, pers. comm.) 
which have a very dark phenotype are regularly encountered on Hooper Hill and in 
the Piute Mountains during late August and September. These represent a later 
brood of virgulti. 


64 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


LIBYTHEIDAE 
95. Libytheana bachmanii larvata (Strecker). 


This is not a regular member of the county fauna. Several individuals of bachmanii 
were observed flying southward through Bakersfield in September and October of 1963. 
I captured one such individual on 6 X 63. Another specimen had been captured two 
years earlier in Bakersfield by Glenn Broadwater. No such migrations have been noted 
since 1963. 


HELICONIIDAE 
96. Agraulis vanillae incarnata (Riley). 
Distribution: SJV, one record for Caliente. Flight: 13 IV to 3 XII. 


NYMPHALIDAE 
97. Speyeria coronis hennei (Gunder). 


Distribution: TM, TJM. Flight: 6 VI to 28 VIII. 

Hennei is often uncommon and hard to find. It prefers the high elevation cooler 
slopes of Mt. Pinos and Double Mountain and rarely descends to lower warmer ele- 
vations. 


98. Speyeria callippe macaria (Edwards). 


Distribution: MTNS. Flight: 14 V to 3 VIII. 
Unsilvered laurina Wright occurs in all the Kern County populations so I consider 
it to be a “form” rather than a valid subspecies. I have one record of macaria from 


Bakersfield, 2 VI 62. 


99. Speyeria egleis tehachapina (Comstock). 


Distribution: Above 7000 feet elevation in TM, PM. Limited to summit peaks and 
ridges in TM. Flight: 30 VI to 24 VII. Records exist for August (Emmel & Emmel, 
1973). 

This endemic subspecies is found only in Kern County. On Double Mountain a forest 
fire consumed much of the summit area in 1979, but adults could still be found after- 
wards, perching on the blackened soil and rocks on the summit. 


100. Speyeria adiaste atossa (Edwards). 


Distribution: TM, TJM. Flight: June to early September. 

Some recent authors (Miller & Brown, 1981; Pyle, 1981) view the adiaste (Edwards) 
group to be subspecies of Speyeria egleis (Behr). However, S. egleis tehachapina and 
S. adiaste atossa were once sympatric in the Tehachapi Mountains. This is strong 
evidence that the two are best viewed as distinct species. 

Though numerous lepidopterists have visited colony sites and habitat regions of 
where atossa was once found, there are still no records of capture since 1959. It has 
probably become extinct because of drought or overgrazing (Emmel & Emmel, 1973; 
Howe, 1975). 


101. Speyeria hydaspe viridicornis (Comstock). 
Distribution: GM. Flight: 2 VI to 3 VIII. 


102. Clossiana epithore sierra (Perkins). 


Distribution: Tiger Flat Campground, GM. Flight: 24 VI to 11 VII. 

This species still flies in the Greenhorn Mountains north of the Tulare County line 
butits status at Tiger Flat Campground is questionable. The author knows of no records 
for the past few years, and several visits to the colony site in June and early July (1981) 
failed to turn up a single specimen. 


103. Thessalia leanira (Felder & Felder). 


At least two distinct populations occur in the county: 


VOLUME 37, NUMBER 1 65 


a. Thessalia leanira nr. wrightii (Edwards). 


Distribution: SNM (Pacific Crest Trail), GM (near Kernville), Erskine Creek nr. 
Lake Isabella, Bodfish, PM, Havilah. Flight: 24 IV to 19 VI. 

This population is extremely variable. About 70% of the population near Bodfish 
is wrightii; 25% resemble daviesi (Wind) or nominate leanira; and the remaining 
5% resemble the brick red desert subspecies, cerrita. 


b. Thessalia leanira cerrita (Wright). 
Distribution: RRC, WP. Flight: 1 IV to 21 V. 


104. Charidryas palla palla (Boisduval). 


Distribution: MTNS, KV. Flight: 1 V to 6 VII. 

Palla is extremely variable from population to population even within the county 
boundaries. In the southern Sierra (Fay Creek 6 miles north of Weldon) and in Kelso 
Valley the palla pattern tends towards obsolescence and individuals may even resem- 
ble C. neumoegeni. Populations to the west have progressively more heavily patterned 
individuals which little resemble the palla found to the east. 

At Kelso Valley the ranges of palla and neumoegeni overlap. There appears to be no 
evidence of any intergradation taking place, however, as the newmoegeni population 
shows no tendency to assume palla characteristics, and the two species are allochronic 
in their occurrence. Thus, neumoegeni flies in desert washes in April and palla in 
those same desert washes in May. I have never found the two species on the wing at 
the same time, though only 4 or 5 days separate the flight periods of the two at this 
locality. 


105. Charidryas neumoegeni neumoegeni (Skinner). 
Distribution: MD, KV. Flight: 5 IV to 7 V. 


106. Charidryas gabbi (Behr). 


Distribution: TJM. Flight: Late May to June. 

The status of this spécies is uncertain. Gabbi is frequently reported from the Frazier 
Park region, but many of these records are based on misidentifications of the heavily 
patterned C. palla which inhabits the area. However, I feel that at least some of these 
records are valid as gabbi is known to be present in nearby areas, and there is apparent 
contact between palla and gabbi at Frazier Park and on the north slope of Frazier 
Mountain. I collected a small series of Charidryas adults in this region (22 VI 79) 
which show startling mixed characteristics of the two species. John F. Emmel examined 
these specimens and concluded that there is obvious gene flow between the two en- 
tities in western Kern County. Further field work and study is needed. 


107. Phyciodes pratensis (Behr). 


Distribution: SNM, GM, KRV, Havilah, Miracle Hot Springs in KRC. Flight: 9 V to 
8 IX. 

Miller and Brown (1981) favor the use of pratensis over that of the more commonly 
used name, campestris (Behr). 

The Kern County population of this species is atypical and may represent an unde- 
scribed subspecies. Most individuals tend towards montana (Behr), but specimens 
which approach nominate pratensis are not unusual (approx. 30-35% of the popula- 
tion). 


108. Phyciodes mylitta mylitta (Edwards). 

Distribution: KRV, MTNS. Flight: 15 III to 24 X. 

A few records of strays exist for the southern San Joaquin Valley. 
109. Occidryas chalcedona chalcedona (Doubleday). 


Distribution: MTNS. Flight: 23 IV to 4 VII. 
Material from the mountains of northeastern Kern County have enlarged light yellow 
spots and tend towards olancha Wright. 


66 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


110. Occidryas editha editha (Boisduval). 


Distribution: WP, SNM, GM (Cedar Creek), PM, KRC (rare). Flight: 20 IV to 8 VII. 
Tends to be very local. 


111. Polygonia satyrus satyrus (Edwards). 


Distribution: MTNS, KRV, SJV (Kern River at Hart Park; Poso Creek 8 miles north 
of Oildale). Flight: 17 III to 4 IX. One record for 19 I 76 at Richbar in KRC. 

Very dark forms resembling neomarsyas dos Passos are frequently encountered in 
the early spring while the lighter chrysoptera Wright is found during the summer 
months. 


112. Polygonia zephyrus (Edwards). 


Distribution: PM, GM, Mt. Pinos. Flight: 12 IV to 17 VII. 
Zephyrus tends to be local in occurrence and less common in Kern County than it 
is in the Sierra further north. It is usually found at higher elevations than P. satyrus. 


113. Nymphalis californica californica (Boisduval). 


Distribution: KRV, MTNS. Flight: 21 II to 11 VII; 4 X to 6 X. 

This species periodically undergoes dramatic fluctuations in numbers. In 1972 and 
1973 N. californica was very abundant. In the drought years 1977 and 1978 I saw none 
at all. It occasionally strays into the southern San Joaquin Valley with records from 
Bakersfield (27 V 52, 4 males leg. Allen Rubbert; 4 X 61; 14 V 72) and the Kern River 
at Hart Park (6 X 71). 

114. Nymphalis antiopa antiopa (Linnaeus). 

Distribution: KRV, SJV, MTNS. Flight: All months. 


115. Aglais milberti furcillata (Say). 


Distribution: KRV, MTNS. Flight: 20 II to 20 VII. 

This species can be very common at times in the Piute, Greenhorn and Tehachapi 
mountain ranges. A few records also exist for the lowlands as it has been collected 
along the Kern River at Hart Park (31 III 72; 4 IV 72) and at Bakersfield (22 IV 51, 2 
males leg. Allen Rubbert). Jim Brock found larvae at Hart Park on nettle (Urtica ho- 
losericea Nutt.) in 1972. These records only prove that furcillata sometimes strays to 
the Valley floor and establishes small transient populations. It is not found at these 
localities in the spring on a yearly basis. 

116. Vanessa virginiensis (Drury). 
Distribution: JC, SJV, KRV, MTNS. Flight: 12 II to 24 X. 


117. Vanessa cardui (Linnaeus). 


Distribution: Entire county. Flight: All months. Has been noted to overwinter on 
low foothills around Hart Park. It is sometimes extremely abundant during migrations. 


118. Vanessa annabella (Field). 

Distribution: Entire county. Flight: All months. 
119. Vanessa atalanta rubria (Fruhstorfer). 

Distribution: SJV, KRV, MTNS. Flight: All months. 
120. Junonia coenia Hubner. 

Distribution: Entire county. Flight: 20 II to 7 XI. 


121. Basilarchia lorquini lorquini (Boisduval). 


Distribution: SJV (Kern River at Hart Park; Poso Creek 8 miles north of Oildale), 
KRV, MTNS. Flight: 9 IV to 24 X. 


VOLUME 37, NUMBER 1 67 


122. Adelpha bredowii californica (Butler). 
Distribution: KRV, MTNS. Flight: 12 IV to 3 XI. 


SATYRIDAE 
123. Coenonympha california california Westwood. 


Distribution: KRV, MTNS. Flight: 25 II to 3 X. 

I collected four specimens of this species along Poso Creek, 8 miles north of Oildale, 
on 4 V 81. These may represent an established population. The species is not generally 
found on the floor of the San Joaquin Valley. 


124. Cercyonis sthenele silvestris (Edwards). 


Distribution: MTNS, WP, KV. Flight: 28 V to 18 IX. 

I follow the prevailing view of recent authors that silvestris is a subspecies of sthe- 
nele (Boisduval) and not Cercyonis oetus (Boisduval) as listed in Miller & Brown 
(1981). 


DANAIDAE 
125. Danaus plexippus (Linnaeus). 


Distribution: Entire county. Flight: All months. 

The Monarch overwinters in moderate numbers along the lower Kern River at Hart 
Park and Lake Ming. It also overwinters in the College Heights residential area of 
Bakersfield. 


126. Danaus gilippus strigosus (Bates). 


Distribution: SJV (rare), KRV, Havilah, Caliente region, KV, MD. One record for 
SNM (Fay Creek, 6 miles north of Weldon; 14 VI 80). Flight: 14 VI to 7 X. Jim Brock 
has one record for April. 

Freezing temperatures in the winter may make it impossible for this butterfly to 
overwinter in the county. Apparently, migrants regularly reach the area in the spring 
and establish breeding populations at various favorable locations. These localities are 
usually in lower mountain canyons or valleys where the narrow-leaved milkweed As- 
lepias fascicularis Dene. in A. DC. is locally common along streams or drainage ditch- 
es. Freshly emerged adults (including one which was deformed and unable to fly) and 
mature larvae of strigosus have been found near or on this plant, suggesting that this 
is the primary larval foodplant for the region. 

Prior to 1981 rare strays had been collected in the southern San Joaquin Valley. 
These records included the following: Hart Park (? IV 68 leg. Jim Brock; 25 VII 70; 
18 VIII 70; 27 IX 70); Bakersfield (11 IX 60; 13 IX 63); 6 miles south of Greenfield 
adjacent to Hwy. 99 and Union Ave. (21 IX 79; 18 IX 81). 

Considering the fact that this species has been so rarely collected in the Valley it 
came as quite a surprise to find a breeding population on the Tule Elk State Reserve 
on 20 VIII 81. I observed no fewer than ten different individuals of strigosus on that 
afternoon. Five were captured. Two additional specimens were collected at the same 
locality on 5 IX 81. Aslepias fascicularis was very common along the wash bottoms 
and swampy region where strigosus was encountered. No other Aslepias species was 
seen on the Reserve. 


Doubtful or Questionable Records 


Nastra julia (Freeman). 

I collected a single specimen from an alfalfa field in Bakersfield during the summer 
of 1962 (no specific date). The specimen was tentatively identified by John F. Emmel. 
We know of no additional records. 

Poanes zabulon (Boisduval & LeConte). 
Emmel & Emmel (1973) reported that there is a “female of this species in the Na- 


68 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


tional Museum of Natural History (USNM) collection with the label “‘Havilah, Calif./ 
July/Barnes Collection.” Like the Emmels, I doubt its authenticity. 


Heraclides cresphontes (Cramer). 


Though recent publications (Emmel & Emmel, 1973; Tyler, 1975) have suggested 
that cresphontes may have extended its range into Kern County and the Central Valley 
of California, we have no records for this species from Kern County. These statements 
have been made on the basis of larvae discovered in Fresno County. This infestation 
has since been eradicated. 


Chalceria rubidus (Behr). 


Emmel & Emmel (1973) report a possible record of this species from Monolith listed 
in the 1950 Field Season Summary. No populations of rubidus are known from this 
region today, and the record is considered doubtful. 


Icaricia shasta (Edwards). 


Emmel & Shields (1978(80)) report that there is a single male in the Los Angeles 
County Museum labelled “Tehachapi Mts., Kern Co., Calif.” collected 22 VIII 37 by 
W. A. Evans. The record is doubtful since it has not been duplicated by others in this 
relatively well collected mountain range. 


ACKNOWLEDGMENTS 


Tule Elk Reserve is in the state park system; so, I am indebted to K. R. Morgan (the 
Area Manager) for providing the necessary collecting permit. 

I would also like to thank all of the previously named individuals who directly or 
indirectly contributed data or information used in this paper. Dr. John M. Burns of the 
Smithsonian Institution clarified problems involving the genus Erynnis. Dr. William 
W. McGuire was consulted regarding the genus Hesperia. 

Jim Brock was my co-worker on this project and has provided considerable infor- 
mation including foodplant records. He has also pointed out a number of taxonomic 
issues. Dr. John F. Emmel provided similar help through his numerous letters and 
helped clarify some of the taxonomic issues which were faced while writing this paper. 
Robert Langston first suggested I write and publish a Kern County checklist and pro- 
vided considerable suggestions and encouragement which were most helpful. Not all 
of these suggestions from these contributors have been followed and the author takes 
full responsibility for any errors or criticisms of this paper. I would like to thank Dr. 
Emmel and Mr. Langston for reviewing this manuscript. 


LITERATURE CITED 


BuRNS, J. M. 1964. Evolution in the skipper butterflies of the genus Erynnis. Univ. 
Calif. Publ. Entomol. 37:1-214. 

Comstock, J. A. 1927. Butterflies of California. Priv. publ. Los Angeles, California. 
334 pp., 63 pls. 

EMMEL, J. F. & O. SHIELDS. 1978(80). The biology of Plebejus (Icaricia) shasta in 
the western United States (Lycaenidae). J. Res. Lepid. 17(2):129-140. 

EMMEL, T.C. & J. F. EMMEL. 1973. The butterflies of southern California. Nat. Hist. 
Mus. of Los Angeles County, Sci. Ser. 26:1-148. | 

MILLER, L. D. & F. M. BROwN. 1981. A catalogue/checklist of the butterflies of 
America north of Mexico. Lepid. Soc. Mem. No. 2. 280 pp. 

HowE, W. H., coordinating editor. 1975. The Butterflies of North America. Double- 
day & Co., Inc., Garden City, New York. 633 pp., 97 pl. 

LANGSTON, R. L. 1969. A review of Glaucopsyche, the silvery blues, in California 
(Lycaenidae). J. Lepid. Soc. 23:149-154. 

OPLER, P. A. 1968. Studies on the Nearctic Euchloe: part 5. Distribution. J. Res. 
Lepid. 7:65-86. 


VOLUME 37, NUMBER 1 69 


1969. Studies on the Nearctic Euchloe: part 6. Systematics of adults. J. Res. 
Lepid. 8:153-168. 

PyLE, R. M. 1981. The Audubon Society Field Guide to North American Butterflies. 
Chanticleer Press Inc., New York. 916 pp. 

SHIELDS, O. 1977. Studies on North American Philotes (Lycaenidae). J. Res. Lepid. 
LGC a= 67. 

TYLER, H. A. 1975. The Swallowtail Butterflies of North America. Naturegraph Pub- 
lishers, Inc., Healdsburg, California. 192 pp., 16 pl. 


Journal of the Lepidopterists’ Society 
37(1), 1983, 69 


BOOK REVIEW 


BUTTERFLIES OF OMAN, by Torben Larsen. 1980. J. Bartholomew and Sons, Edin- 
burgh. 80 pp., ill. 


This book is part of a concerted effort by the Omani government to raise the conscous- 
ness and appreciation of that country’s flora and fauna. It succeeds in its mission, both 
scientifically and artistically, the latter apparently largely through the efforts of Larsen’s 
wife, Kiki. 

It will surprise many that there is a significant butterfly fauna in this desert realm, 
but in fact, more species occur there than are found in the British Isles. Most of them 
occur in various wadis, around oases and in urban environments where the vegetation 
is relatively lush; many of the intervening arid areas are basically sterile. The Omani 
fauna is derived from three basic sources: the Palearctic, the Indian region, and arid 
eastern and southern Africa. Examples of each are given in the text. Whatever is known 
of the biology of all of the species is given, along with photographs of foodplants in 
many instances. 

The nomenclature, though it will not please “traditionalists,” is up-to-date and in 
conformity with that employed in the Palearctic literature. Thus, some “‘old friends” 
are in unfamiliar genera: Stonehamia, Artogeia, Pontia, Epamera and Pseudophilotes 
are used for species formerly placed in Charaxes, Pieris, Pieris, Iolaus and Philotes, 
respectively. These generic changes, though, are based on solid biological and mor- 
phological studies, so their acceptance is made easier, even though there will be the 
inevitable complaints. 

One aspect of the nomenclature that I do question involves two “species pairs” 
recognized in the book. One of these is Zizeeria knysna (Trimen) and Z. karsandra 
(Moore), usually considered conspecific. These two entities are both in Oman but at 
opposite ends of the country, apparently not sympatric at all, even in Oman. Since Z. 
knysna is an African entity, and Z. karsandra is Oriental, and both occur about where 
one would expect to find African and Asian faunal elements, their allopatric occurrence 
in Oman does not make a convincing case for their specific identities. The same ob- 
jections can be made for the Asian Papilio demoleus and the African P. demodocus. In 
both instances, the African elements are known only from near Dhofur. 

Despite such nit-picking criticisms as the above, I can recommend the book for 
anyone interested in the butterflies of this area, or even for the reader who is interested 
in what butterflies might be hardy enough to withstand the vicissitudes of such a 
climate. Mr. Larsen, perhaps our premier authority on Middle East butterflies, is to be 
congratulated on another fine book, though perhaps with less scientific “meat” than 
his earlier Butterflies of Lebanon. 


LEE D. MILLER, Allyn Museum of Entomology, 3701 Bay Shore Road, Sarasota, 
Florida 33580. 


Journal of the Lepidopterists’ Society 
37(1), 1983, 70-77 


ERBLICHIA ODORATA SEEM. (TURNERACEAE) IS A 
LARVAL HOST PLANT OF EUEIDES PROCULA VULGIFORMIS 
(NYMPHALIDAE: HELICONIINI) IN SANTA ROSA 
NATIONAL PARK, COSTA RICA 


D. H. JANZEN 


Department of Biology, University of Pennsylvania, 
Philadelphia, Pennsylvania 19104 


ABSTRACT. The larva of Eueides procula vulgiformis Butler & Druce feeds on 
the leaves of Erblichia odorata, a large turneraceous evergreen tree that occurs in 
relatively pristine evergreen forest patches among the deciduous forest patches of Santa 
Rosa National Park (300-350 m elevation) in the northwestem coastal plain of the 
Pacific side of Costa Rica. This is the only known case of a heliconiine caterpillar 
feeding on the foliage of a tree and the only known case of a heliconiine caterpillar 
feeding on a plant outside of the Passifloraceae. The larval and pupal stages are de- 
scribed and figured and are unexceptional for the genus Eueides. 


Eueides procula vulgiformis Butler & Druce is a common butterfly 
(Fig. 1) in the small patch of evergreen forest (“Bosque Humedo,”’ 
“Bosque Siempre Verde’) along the main entrance road in the north- 
east end of Santa Rosa National Park, northeastern Guanacaste Prov- 
ince, Costa Rica (350 m elevation). Its larval host and larval stages 
were unknown until now (K. Brown, 1981; P. J. DeVries, pers. oui 
and are described here. 

Adults are present in the Bosque Himedo throughout the year, but 
fluctuate strongly in density within and between years. From Decem- 
ber 1979 through January 1982, they have been abundant in a forest 
where they were not encountered during intensive collecting in 1978 
and early 1979 by P. J. DeVries, L. E. Gilbert and J. J. Smiley. In 
December 1979 and later, an adult could be located with a few min- 
utes search. The most common and omnipresent heliconiine in this 
habitat is Heliconius hecale zuleika Hewitson, with which E. p. vul- 
giformis is a very good Miullerian co-mimic. Adult E. p. vulgiformis 
are encountered throughout the daylight hours, flying and soaring 
from ground level to the tops of the tallest trees (20 m); when 10-20 
m tall Licania arborea (Chrysobalanaceae) are in flower in late Decem- 
ber and early January, members of both sexes are common visiting 
the flowers, and males chasing other males are a commonplace. 

The larval host of E. p. vulgiformis in the Bosque Humedo is a 
large turneraceous tree, Erblichia odorata Seem. The tree is common 
in this habitat, attains a height of 15-20 m (DBH up to 50 cm), and is 
evergreen. E. odorata does not occur in the deciduous forests sur- 
rounding the Bosque Humedo and E. p. vulgiformis is not encoun- 
tered in these forests either. E. odorata has simple and glabrous lan- 


VOLUME 37, NUMBER 1| valk 


Fic. 1. Eueides procula vulgiformis adults reared from larvae found on Erblichia 
odorata in Santa Rosa National Park, Guanacaste Province, Costa Rica: Upper left, 
female; Lower left, female underside; Right, male. 


ceolate leaves 8-14 cm long and 1-3 cm wide (Fig. 2a), with a gently 
undulating and slightly toothed margin. The leaves are less stiff and 
leathery than those of most of the evergreen tree species in the Bos- 
que Humido but somewhat more stiff and leathery than those of most 
of the deciduous trees in the area. While most of the individuals of 
E. odorata are adult or subadult trees, the forest is sprinkled with a 
small number of saplings and seedlings. 

When the density of E. p. vulgiformis is high, females are often 
seen fluttering about the margins of the crowns of large E. odorata, 
both in sunlit margins of tree-falls and in moderately shady portions 
of a well-closed canopy. Males chase them at this time, and sometimes 
the advances of a male stops an ovipositing female from laying an egg 
and causes her to move on. If a female is ovipositing on leaves in a 
small E. odorata sapling intermingled with foliage of other trees, she 
often lands many times on leaves of other species before landing again 
on an E. odorata leaf. For example, during an hour of fluttering about 
in the foliage and avoiding advances by males, one female E. p. vul- 
giformis contacted eight E. odorata leaves and laid a single egg on 
each of four of them. She landed on 37 leaves of other species during 
this period (1000-1100 h, 2 July 1980; weather was sunny and breezy). 
She always landed on the upper surface of the E. odorata leaves, and 
the weight of her body caused the leaf to bend sharply downward. 
She then reached under the leaf with her abdomen and glued a single 
egg to the central portion of the blade, 1-2 mm to one side of the 


op) JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


ere 


agra ioc pce eT 


nye piracy peer eT TA Vth} 


STITT 


Fic. 2. a, normal-sized mature leaf of Erblichia odorata with a single egg of Eueides 
procula vulgiformis laid on its underside slightly to the left of center; b, egg of E. p. 
vulgiformis; e, second instar larva of E. p. vulgiformis feeding at the margin of the 
characteristic holes they cut in the leaf in the first and second instar. All scales in mm 
in this and later figures (all photos in this and later figures, Santa Rosa National Park, 
Guanacaste Province, Costa Rica, 350 m elevation). 


midrib (Fig. 2a). Immediately after laying the egg she launched into 
a fluttering flight among the foliage; even if there was another E. 
odorata leaf nearby, she showed no directed flight toward it. After 
flying 1-3 m, she again alighted. If the landing site was not an E. 
odorata leaf, she launched into flight in a few seconds, and flew another 
few cm to am or a bit more before landing again. Her general flight 
trajectory was neither straight nor strictly horizontal but rather wan- 
dering in both a vertical and horizontal plane. When chased by a male, 
she sometimes flew as much as 10 m before landing again. When the 
sun was obscured by a cloud for a few minutes, she landed on the 
upper surface of a leaf and sat motionless. | 

The eggs are pale greenish-yellow and have the appearance of 
domed squat cylinders (Fig. 2b). There are numerous ridges running 
from the base to top (portion away from the leaf), with horizontal 


VOLUME 37, NUMBER l| 13 


troughs breaking the ridges into a series of bumps. They look very 
similar to the eggs of Eueides tales figured by Brown (1981), except 
that where the E. tales egg has depressions on the surface, the E. p. 
vulgiformis egg has domes or bumps. 

Eggs were laid on leaves of all ages, but in searching for eggs, I 
found more per leaf on vertical shoots off main trunks than on leaves 
on branches well out into the margins of the crown. For example, I 
watched one female lay three eggs in 12 minutes on a 30 cm long 
sucker shoot with 12 leaves at 4 m off the ground. Eggs were found 
on the leaves of plants as small as 1 m tall in heavily shaded under- 
story and on leaves in the margins of the crowns of trees 15 m tall 
(obtained by climbing trees and cutting down branches). I saw fe- 
males ovipositing in the foliage of the crowns of the tallest trees. 

Of a set of four eggs laid by one female on 2 July, one hatched on 
5 July and the other three on 6 July. The larvae were maintained at 
room temperature (not very different from that in the Bosque Humedo 
3 km away) in large plastic bags and pupated on 24—26 July. Through- 
out the 20 day larval period, the caterpillars were given freshly cut 
E. odorata leafy branches every other day. Still maintained at room 
temperature, each pupa produced a normal-sized adult (3 males, 1 
female) eight days after pupation. 

The first instar larva begins feeding in a very distinctive manner, 
and the second instar continues in the same manner. It eats a ragged- 
edged hole from 0.1 to 0.3 cm? in area (Fig. 2c), through the leaf blade 
within about 5 mm of where the egg was laid, and then moves on and 
eats other similar holes in the same or nearby leaves. After molting 
to the third instar, the larva begins eating at the margin of the leaf 
tip. It consumes the blade down to the midrib on one side and may 
continue until a quarter or more of the leaf blade is missing. Even 
when many caterpillars were confined in the same rearing container 
on a small amount of foliage, there was no sign of gregarious or side- 
by-side feeding. 

The second instar larva (Fig. 2c) is semi-translucent light green with 
two rows of massive spines dorsally and along each side. The spines 
are as long as the body is thick and have spinelets projecting out of 
them. The spines are nearly black and their contact point with the 
body wall is in the center of a whitish-blue ring (dorsal rows of spines 
only). This gives the impression that the caterpillar has a light gray 
stripe down its back. The head capsule bears dorsally a pair of wide- 
spread massive spines like those of the body, but they are curved 
slightly backwards. The spines on the last two segments are paler in 
color than are those on the remainder of the body. The head capsule 
is grayish-brown in color. 


74 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


- te €:, 


Fic. 3. a, third instar larva of E. p. vulgiformis, lateral view; b, third instar larva 
of E. p. vulgiformis, dorsal view. 


The third instar larva gives the overall impression of being white 
with black spots and a yellow posterior (Fig. 3). The main body spines 
are black, as long as the body is wide, and bear conspicuous lateral 
spinelets. The spines do not urticate in this or any other instar. The 
head capsule spines are white, black-tipped, and curve backwards. 
The lateral spines on the posterior two segments are gray-white, and 


VOLUME 37, NUMBER Il VS 


Fic. 4. a, fifth instar larva of E. p. vulgiformis shortly after molting from the fourth 
instar; b, fifth instar larva of E. p. vulgiformis after it has stopped feeding and begun 
spinning the pad on which it will pupate; ec, pre-pupa of E. p. vulgiformis. 


76 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Fic. 5. Pupa of E. p. vulgiformis. 


the ventral lateral spines along the entire body are more gray than 
black. The head capsule is black in color. Dorsally, each body seg- 
ment is white with seven black dots, except that the penultimate seg- 
ment is bright yellow dorsally. Laterally the caterpillar has a brown- 
ish-white stripe bordered by a white stripe ventrally. 

The last (fifth) instar larva, while still feeding, is gray-white with 
a lateral pale yellow stripe (each spiracle covered by a black dot) and 
with a dorsal band of black dots (Figs. 4a, b). All spines are pure 
black, have lateral projections, and are as long as the body is thick. 


VOLUME 37, NUMBER 1 Hol 


The head capsule is black with white markings. The penultimate body 
segment is dorsally orange-yellow. Shortly before it becomes a non- 
feeding pre-pupa, the caterpillar is 22-25 mm in length. 

The color change to the pre-pupa is dramatic. The pre-pupa has a 
bright yellow body with a white head capsule and black spines, ex- 
cept that the most lateral (ventral) set of body spines is white, and the 
base of each body spine is heavily ringed in black (Fig. 4c). The pre- 
pupa wanders for a few hours on the walls of the rearing container 
and then spins a silk pad as a pupation site, or it may change color 
after spinning the silk pad. 

The pupa is white and sparsely flecked with small black specks, 
bears a pair of white but black-tipped spines ventrally on each of three 
abdominal segments, and bears a pair of long slightly hooked white 
spines with gray and yellow tips at the head end (Fig. 5). 

In the forest I have found pupae on leaves of E. odorata and on 
leaves up to a few m from foliage of E. odorata. 

In short, the immature stages of E. p. vulgiformis are in no mor- 
phological way exceptional in comparison with those of other heli- 
coniines (Brown, 1981) but are distinguishable from those of other 
Eueides (K. Brown, pers. comm.). However, the larvae of E. p. vul- 
giformis are quite exceptional in their choice of food plants. This is 
the first unambiguous record of a heliconiine feeding on a plant out- 
side of the Passifloraceae (K. Brown, pers. comm.). However, the 
species has not strayed far, since Turneraceae is closely related to 
Passifloraceae and Turneraceae is a well known host family for other 
Nymphalidae that are closely related to Heliconiinae (K. Brown, pers. 
comm.). 


ACKNOWLEDGMENTS 


This study was supported by NSF DEB 80-11558 and by Servicio de Parques Na- 
cionales de Costa Rica in general and Parque Nacional Santa Rosa specifically. P. J. 
DeVries alerted me to take special note of the butterfly, and he identified the adults, 
and K. S. Brown and P. J. DeVries offered constructive commentary on the manuscript. 
W. Hallwachs found the first pupa of E. p. vulgiformis on E. odorata. 


LITERATURE CITED 


Brown, K. S., JR. 1981. The biology of Heliconius and related genera. Ann. Rev. 
Entomol. 26:427-456. 


Journal of the Lepidopterists’ Society 
37(1), 1983, 78-79 


GENERAL NOTES 


THE SUITABILITY OF JUNIPERUS (CUPRESSACEAE) FOR LARVAE OF 
CALLOPHRYS HESSELI (RAWSON AND ZIEGLER) (LYCAENIDAE) 


Aside from the wing character differences and distinct male and female genitalia 
distinguishing Callophrys hesseli from other Nearctic Cupressaceae-utilizing Callo- 
phrys (Mitoura) (Rawson & Ziegler, 1950, J. N.Y. Entomol. Soc. 58:69-82; Johnson, 
1978, J. Lepid. Soc. 32:3-19, and 1976, Bull. Allyn Mus. 38:1-30), its utilization of 
Chamaecyparis thyoides (L.) B.S.P. as the larval foodplant is a major taxonomic trait 
distinguishing it from its nearest spatial relative C. (Mitoura) gryneus Hubner. The 
latter is known to feed on four species of Juniperus L. (Johnson, 1978, loc. cit.). The 
two butterfly species are regionally sympatric but allopatric on the local level, except 
for occasional overlap in usage of nectar sources (Johnson, 1978, loc. cit.). No naturally 
occurring hybrids or instances of larvae feeding on the foodplant of the other species 
are presently known. 

In laboratory studies, Remington & Pease (1955, Lepid. News 9:4—6) demonstrated 
larvae of gryneus could be raised to the imago stage on C. thyoides and not be nutri- 
tionally sterile. In their conclusions concerning suitability of C. thyoides for gryneus 
they stated the importance of knowing whether Juniperus virginiana L. (the foodplant 
of gryneus in eastern North America) was equally suitable in the laboratory for hesseli. 

During studies of Nearctic Callophrys (Mitoura) (Johnson, 1984, Bull. Am. Mus. Nat. 
Hist., in press) this author received a number of second-hand reports of rearing of 
hesseli on J. virginiana; however, none could be verified with accurate data. The 
purpose of this note is to report the first verifiable rearing of hesseli on Juniperus. 

Eric Quinter (American Museum of Natural History, New York) collected a series of 
live hesseli females on 17 May 1971, 3.6 mi. E of Chatsworth, Burlington Co., New 
Jersey. All were confined under incandescent light in mesh bags above sprigs of C. 
thyoides (in anticipation of rearing all larvae on this plant). Oviposition soon took place 
and over a hundred larvae were reared on C. thyoides until all larvae were in the 
second instar. When available foodplant supplies dwindled about 50 larvae in the 
second instar were transferred to Juniperus virginiana in hopes they might survive. 
All continued to feed readily and maintain normal growth. These larvae reached the 
last instar showing no difference from those on C. thyoides. Then a fungal infestation 
attacked the entire rearing apparatus, occurring just prior to expected pupation and 
resulting in a high mortality rate in larvae on both foodplants. In fact, of the healthy 
Jarvae on J. virginiana, only one last instar larva escaped infestation; however, it pu- 
pated readily and emerged on 10 July. Unfortunately, it was impossible to mate this 
male individual with a reared female from the same foodplant to test possible sterility. 

This rearing demonstrates that J. virginiana is at least suitable for adult maturation 
in Callophrys hesseli. It is particularly impressive because the transference of larvae 
in middle or later instars is usually considered highly unfavorable to normal maturation 
or survival (Downey & Dunn, 1965, Ecology 45:172-178; Dethier, 1954, Evolution 8: 
33-54). Usually in rearing experiments either ova or freshly emerged larvae are trans- 
ferred to an alternative foodplant. Suitability of J. virginiana after the first instar, how- 
ever, still does not prove full laboratory compatibility. Mortality might have occurred 
during the first instar itself. Also, with lack of testing for nutritional sterility, and more 
importantly, oviposition preference of reared females, this rearing can only provide 
limited conclusions. Any adequate test of ovipositional specificity in reared females 
would require larvae reared solely on one or the other plant. However, it is significant 
to record an instance of successful rearing to the adult stage on the alternative butterfly/ 
foodplant combination used by Remington & Pease. Although suitability in the labo- 
ratory is no test of suitability in nature (Downey, 1962, System. Zool. 11:150-159; 
Downey & Dunn, loc. cit.; Dethier, loc. cit.), it is further evidence that specificity in 
the natural environment (which in the case of C. hesseli and C. gryneus seems justi- 


VOLUME 37, NUMBER | 79 


fiably assumed by the abundant field data available on these species) is preserved by 
the oviposition habits of the female, according to Hopkins’ Hostplant Principle. 


KuRT JOHNSON, Dept. of Entomology, American Museum of Natural History, Cen- 
tral Park West at 79th St., New York, New York 10024. 


Journal of the Lepidopterists’ Society 
37(1), 1983, 79-80 


THE TYPE OF ARGYNNIS APACHEANA SKINNER 


In 1954 (Trans. Amer. Entomol. Soc. 80:91-117) Gillham & Ehrlich published an 
excellent review of the butterfly names established by Henry Skinner. They questioned 
the statement in dos Passos & Grey (1947, Amer. Mus. Novitates, No. 1370, 30 pp.) 
that the type of Argynnis apacheana Skinner is from Arizona and in the Academy of 
Sciences, Philadelphia. The specimen referred to by dos Passos & Grey was in the 
collections of the Academy, but has been transferred to the Carnegie Museum of Nat- 
ural History, Pittsburgh, Pennsylvania. It is designated ANSP No. 7031 and carries 
labels reading “holotype” and “Arizona, collector Skinner.” This is contrary to Skin- 
ners declaration of the type of apacheana in the original description, quoted by Gill- 
ham & Ehrlich. 

Skinner (Entomol. News 29:67, lst paragraph) wrote: “I propose the name apacheana 
for the species of Argynnis described and figured by Mr. W. H. Edwards in Volume I 
of his Butterflies of North America, plate IV of Argynnis, figures 1, 2, 6, 3, 4, 2, under 
the name nokomis.” There is no other declaration of type in the article. The butterfly 
figured on this plate is not the type of the name nokomis. In fact, the plate in bound 
copies of the volume is not the original plate which was drawn from the type by Wiest. 
The plate referred to is the replacement for that plate. The new plate was drawn by 
Mrs. Mary Peart from specimens collected by the Wheeler Expedition in 1871, eight 
years after nokomis had originally been described. 

Almost everyone has been misled by the locality designation “Arizona” for material 
sent east by the 1871 Wheeler Expedition. This designation is very much like the old 
one “Bogota” for Colombian butterflies; meaningless. A timetable and route for the 
expedition was published by Brown in 1957 (J. N.Y. Entomol. Soc. 65:219-234). The 
cases of specimens for the Smithsonian Institution were dispatched from Tucson, Ar- 
izona, the breakup point, in December 1871. These boxes contained material from 
most of the route. All that Baird told Edwards when he transmitted the material is that 
he had received it from Arizona (Brown, F. M., 1965, Trans. Amer. Entomol. Soc. 91: 
J35=3 50) 

Some years ago I sent Scott Ellis and Samuel Johnson to Owens Valley, California 
to see if they could recover the species collected there by Bischoff and thus, narrow 
down type localities. They were successful in general but unsuccessful for Cercyonis 
wheeleri and Speyeria “nokomis” (apacheana). Diversion of water from Owens Valley 
by the city of Los Angeles has destroyed the niches in which these sensitive species 
had lived. It is only in the vicinity of Round Lake, where the requisite Viola grows in 
the understory of the meadow and bog grasses, that apacheana is still found. 

Dr. dos Passos is very ill, so I asked Mr. Paul Grey to correct the type statement, but 
he asked that I do it. Since the figure upon which Skinner based his name apacheana 
was drawn from material collected by Bischoff in 1871, that material must supply the 
type. The specimen that was used by Mrs. Mary Peart for the model of the male figure 
on plate Argynnis IV must be considered the type of apacheana. That specimen was 


80 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


retained by W. H. Edwards and now is with his collection in the Carnegie Museum of 
Natural History, Pittsburgh, Pa. Its locality of capture at this time can be given no more 
closely than vicinity of Independence, Inyo Co., California, from a colony now extinct. 


F. MARTIN BROWN, Wright-Ingraham Institute, Colorado Springs, Colorado 80911. 


Journal of the Lepidopterists’ Society 
37(1), 1983, 80-81 


A NEW FOOD PLANT RECORD FOR CHLOSYNE GORGONE CARLOTA 
(REAKIRT) (NYMPHALIDAE) 


Host plants recorded for the larval stages of Chlosyne gorgone carlota (Reakirt) in- 
clude a wide variety of genera in several families. While most authors (Klots, 1951, 
Field Guide to the Butterflies, Houghton Mifflin Co.; Forbes, 1960, Lepidoptera of 
New York and Neighboring States, Part IV, Cornell Univ. Agr. Expt. Sta. Memoir 371; 
Ehrlich & Ehrlich, 1961, How to Know the Butterflies, Wm. C. Brown Co.; Tietz, 1972, 
An Index to the Described Life Histories, Early Stages, and Hosts of the Macrolepi- 
doptera of the Continental United States and Canada, Allyn Mus. Entomol., Sarasota, 
FL; Johnson, 1972 (1973), J. Res. Lepid. 11(1):1-64) list Aster spp. and Helianthus 
spp. (Compositae) as the primary food plants, C. g. carlota larvae have also been re- 


Fics. 14. Female specimens of Chlosyne gorgone carlota from Missouri: 1, 2, 
female showing typical markings of this species, dorsal and ventral views, respectively; 
3,4, female reared from Ambrosia trifida, dorsal and ventral views, respectively. 


VOLUME 37, NUMBER 1 81 


corded from Lippia lanceolata and L. nodiflora (Verbenaceae) (Kimball, 1965, Arthro- 
pods of Florida and Neighboring Land Areas, Vol. 1, Lepidoptera of Florida, Florida 
Dept. Ag.), Lysimachia sp. (Primulaceae) (Harris, 1972, Butterflies of Georgia, Univ. 
of Oklahoma Press), and Eriogonum sp. (Polygonaceae) (Tietz, ibid.). - 

In Missouri, C. g. carlota larvae have been recorded from Helianthus annuus L. 
(Compositae) (J. R. Heitzman, pers. comm.) and Linaria vulgaria Hill (Scrophularia- 
ceae) (Masters, 1969, J. Kansas Entomol. Soc. 42(2):133-144). Here I report giant rag- 
weed, Ambrosia trifida L. (Compositae), as a new food plant for C. g. carlota. A. trifida 
L. is an annual, monoecious weed common to fertile moist soils, bottom lands, allu- 
vium, and waste places. 

On 9 August 1977, while collecting along a roadside bank of State Rt. 10 (3.2 km 
west of Richmond, MO), I found thirteen larvae that were unfamiliar to me on the 
leaves of three separate giant ragweed plants. Due to the uniformity in larval size and 
the close proximity of the plants to each other (within 0.5 m of each other), the infes- 
tation was probably the result of a single oviposition. The thirteen larvae, along with 
an ample supply of the food plant, were collected and taken to my home to be reared. 
Six larvae were preserved on 9 August 1977; the remaining seven larvae were allowed 
to feed. On 10 August 1977 one larva pupated, emerging 11 days later. The single adult, 
identified as a female C. g. carlota by J. R. Heitzman, is slightly smaller and darker 
than the typical female of the species (Figs. 1-4), which may have resulted from the 
rearing. The six remaining larvae pupated on 11 August 1977 and were preserved. 

I am especially grateful to J. Richard Heitzman for identifying the specimen and for 
reviewing this manuscript. Thanks are also extended to Dr. Charles V. Covell, Jr., Dr. 
Stephen Clement and to Ms. Candy Fogg for their critical reviews and to Mr. Glenn 
Berkey of the Ohio Agricultural Research and Development Center, Wooster, for the 
photographs. 


DAvID C. IFTNER, 2161 Heatherfield Avenue, Worthington, Ohio 43085. 


Journal of the Lepidopterists’ Society 
37(1), 1983, 81-82 


A NATURAL OCCURRENCE OF INTER-TRIBAL 
COPULATION IN THE PAPILIONIDAE 


Among the Lepidoptera numerous prezygotic isolating mechanisms (Mayr, 1970, 
Population, Species and Evolution, Belknap Press, Cambridge, MA) operate to prevent 
interspecific mating. At times, however, these isolating mechanisms appear to break 
down. While breakdowns occur rarely between closely related species, breakdowns 
between distantly related and phenotypically distinct species are quite exceptional. A 
salient example of such a breakdown was observed on 15 April 1980 in the Kirby State 
Forest near Kountze, Tyler County, Texas. 

Swallowtail diversity in Kirby Forest and throughout the upland savannah of south- 
east Texas is high; at least six different species are resident (Rausher, in preparation). 
Eurytides marcellus (Papilionidae: Graphiini) and Battus philenor (Papilionidae: Troi- 
dini) fly sympatrically in the open pine uplands. The two species are temporally syn- 
chronous, adults flying commonly between mid-March and mid-April. During this pe- 
riod females spend much of their time searching among the herbaceous vegetation for 
larval food plants. For B. philenor, these are Aristolochia reticulata and A. serpentaria, 
small erect perennial herbs in the family Aristolochiaceae (Rausher, 1978, Science 200: 
1071-1073). The larval food plant of E. marcellus, in contrast, is Asimina parviflora, 
an annonaceous shrub that grows in east Texas to a maximum height of 2-3 ft. Males 
of each species fly through the pine uplands, approach females while they are ovipos- 


82 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Fic. 1. Female Battus philenor and male Eurytides marcellus photographed in 
copulo in Kirby Forest, southeast Texas. 


iting or nectaring, and engage in precopulatory courtship flights. Although males of 
either species may approach heterospecific individuals, such encounters are usually 
brief in duration. In seven years of observation we have never seen a male engage in 
persistent courtship of a heterospecific female, although we have observed hundreds 
of conspecific courtships. 

A female B. philenor and male E. marcellus (Fig. 1) were found resting in copulo 
among the forest herbs in Kirby Forest. After the pair had disengaged, the female was 
captured and offered foliage of both species of Aristolochia on which to oviposit. She 
failed to lay eggs on either species over a period of three days. Subsequent dissection 
of the female revealed no spermatophore in the bursa copulatrix, although many de- 
veloped chorionated eggs were present. These observations suggest that, despite the 
fact that the two individuals remained in copulo for more than 30 minutes, no sperm 
transfer occurred. 

Since little is known about the courtship and mating behavior of either butterfly 
species, it is difficult even to speculate about the events that gave rise to this unusual 
pairing. Nevertheless, despite the apparent breakdown of ethological isolating mech- 
anisms in this instance, reproductive isolation was maintained by failure of the male 
to transfer sperm. Whether this failure was due to active interference by the female or 
simply to the absence of an appropriate cue necessary for triggering sperm transfer is 
not known. 

We thank M. Nijhout for dissecting the female butterfly and Paul Feeny for bringing 
the east Texas habitat and swallowtails to our attention. This work was supported in 
part by N.S.F. grant DEB 8016414 to MDR. | 


MARK D. RAUSHER, Department of Zoology, Duke University, Durham, North Car- 
olina 27706 AND M. BERENBAUM, Department of Entomology, 320 Morrill Hall, Uni- 
versity of Illinois, 505 South Goodwin, Urbana, Illinois 61801. 


VOLUME 37, NUMBER 1 83 


Journal of the Lepidopterists’ Society 
37(1), 1983, 83-85 


NOTES ON THE COURTSHIP OF TROIDES OBLONGOMACULATUS PAPUENSIS 
(PAPILIONIDAE) IN PAPUA NEW GUINEA 


Straatman (1976, Trans. Lepid. Soc. Japan 27:156—162), gave an account of the court- 
ship behavior exhibited during hybridization of Troides oblongomaculatus papuensis 
Wallace males with Ornithoptera priamus poseidon Doubleday females. However, 
there has been no detailed description of the courtship behavior of T. 0. papuensis, 
particularly the use of the hindwing pouches of the male, although various authors 
have commented upon their highly specialized morphology and obvious androconial 
function. Similar pouches are found at the base of the costa of the forewings of Chae- 
tocneme (Hesperiidae) males from the same region. 

The cultivated Aristolochia tagala Chan. vines in the grounds of the Insect Farming 
and Trading Agency (I.F.T.A.) at Bulolo in Papua New Guinea have produced a thriv- 
ing population of T. 0. papuensis that can be observed readily throughout the year. 
These notes are a summary of observations of over fifty courtships and of a mark- 
recapture study over a period of one year (March ’79—March ’80). 

Mark-recapture results by the author of ex-pupa specimens have shown that individ- 
ual males can live up to two and one-half months and probably longer. They regularly 
patrol the grounds of the I.F.T.A. in search of freshly eclosed females by flying to 
inspect each vine in turn. Up to six males have been in sight at one time and often one 
or two females may be observed ovipositing at the same time. Marked females have 
been recaptured while ovipositing on the vines only up to about a week later, which 
suggests that they go much further afield in search of suitable vines on which to ovi- 
posit; thereby, maximizing the dispersal of the species. 

Numbers of individuals in the I.F.T.A. grounds have always been too low to obtain 
an accurate estimate of population size using standard formulae, but a constantly re- 
newed population of 15-25 individuals would be a fair estimate. The farthest distance 
from which a marked male has been recovered is 2 km, but they probably range farther 
and have been observed to revisit areas on a rotational basis to feed and search for 
females. 

There are two distinct patterns of courtship by Troides males. In the first case, where 
a newly eclosed virgin female is encountered, pairing is abrupt with apparently no 
signal from the female that she is receptive. She merely does not prevent the male 
from coupling. The display by the male prior to pairing is, therefore, short and consists 
of a period of no more than 30 seconds, while he flutters with rapid wing beats close 
to, but not actually touching, the female. The male will settle near her on foliage and 
then turn to engage his open claspers to each side of her abdomen. The female opens 
her genital aperture by raising the ovipositor, and the union is effected within a matter 
of a few seconds. If the female flies off later the male is carried hanging inertly below 
with legs folded. When the female alights again the male will often remain in this 
posture. Based on ten observations, pairing lasts for about five hours. 

The second category of courtship behavior is a sustained and repeated display ini- 
tiated by the sighting of a previously paired, flying female. Males seem unable to 
recognize mated females, and prolonged courtship is only exhibited to such individ- 
uals. The author has never witnessed sustained aerial courtship of virgin females, 
probably because males in the grounds of the I.F.T.A. are always sufficiently numerous 
to locate most new females before they take to the air. Males may even be able to learn 
in advance the position of female larvae and pupae, because on many occasions Troides 
males have been observed to fly and investigate these early stages. Often they will 
hinder ovipositing females or (rarely) pursue other males in a briefly attempted court- 
ship display. 

The display to the female is a fluttering flight for the purpose of engaging her an- 
tennae with the androconial hair-scales of the scent pouches of the male. Therefore, if 


84 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Fic. 1. Left side of head of female Troides oblongomaculatus papuensis, showing 
androconial scales from male adhering to her antenna. 


the female is flying along in a straight line, the male will execute loops around her, 
coming quickly from behind and underneath so that the last wingbeat carries him 
upwards and backwards. The hindwing pouches are thereby timed to open as the 
female moves forward, so that her antennae then engage in the open slots of the male's 
pouches. 

Sustained courtship by the male in this way elicits an avoidance reaction by the 
previously paired female, who then goes to ground with wings spread and abdomen 
pointing downwards. Often the male will continue this display and hover just above 
the head of the female, repeatedly backing his scent pouches in irregular sweeps onto 
the female’s antennae by quick, downward strokes of the forewings. The male may fly 
off and return three or four more times but soon leaves the female at this stage. She 
may remain inert and prostrate on the ground for some minutes after the male has gone 
and then suddenly take to the air. 

If the androconial pouch of the male is opened it can be seen to be full of vertically 
arranged, extremely close packed, white, hair-like scales. These are very easily dis- 
lodged and are so fine and light that they easily stick to a shiny needle point. Their 
function in courtship is to adhere to the antennae of the female (Fig. 1). Therefore, it 
appears that the male’s courtship pheromone is not airborne but is physically trans- ~ 
ferred via the androconial hairs to the chemosensory surface of the female's antennae. 
The photograph shows the head of a female immediately after the repeated courtship 
display of a male. The hair-scales even adhere to the proboscis when the female has 
been subjected to a long courtship display. 

Similar courtship has been observed in Ornithoptera, but it appears that the stiff 
fringe of brush-like hairs on the inner margin of the hindwing of Ornithoptera males 


VOLUME 37, NUMBER 1 85 


is a modification of the scent pouch of Troides. These long hairs are covered with a 
pheromone which is brushed by direct contact onto the antennae of the female during 
the courtship flight, but the hairs are not displaced. If the hindwings of a fresh male 
of O. priamus poseidon, for example, are placed on white card and stuck tightly beneath 
clear sticky-backed plastic, from the scent hairs only an orange compound, which pre- 
sumably contains the pheromone, is slowly leached off through the glue reminiscent 
of a chromatogram. 

During copulation a gelatinous substance is produced by the accessory glands of the 
male. This is soft and clear at first with a slight yellowish tinge and almost fills the 
genital cavity of the female once the male has parted. Later it dries hard and becomes 
opaque and dark brown. This is the sphragis, which is thought to act as a barrier to 
further insemination. However, as some ovipositing females have been found to have 
lost this, it appears more likely that it is the presence of the large spermatophore (which 
fills the bursa copulatrix) that produces a stimulus to reject further males. Nevertheless 
in Cressida and Parnassius (Papilionidae) and Miyana (Nymphalidae) the sphragis is 
external, very large, and is permanent and surely must physically prevent further pair- 
ing. Of the photographs in figure 65 (p. 87) of Haugum & Low (1979, A Monograph of 
the Birdwing Butterflies. Vol. 1, part 2, Scand. Sci. Press) of supposed sphragis in O. 
priamus, only the central picture shows this. The outer two merely figure the artificially 
distended genital plates of the female. Specimens killed by injection with ethy] acetate, 
for example, often die in this latter condition. 


MICHAEL PARSONS, Lepidoptera Research Project, Insect Farming and Trading 
Agency, Division of Wildlife, P.O. Box 129, Bulolo, Morobe Province, Papua New Guinea. 


Journal of the Lepidopterists’ Society 
37(1), 1983, 85-86 


ALBINIC VARIANTS OF CHLOSYNE NYCTEIS FROM 
CONNECTICUT (NYMPHALIDAE) 


On 8 July 1979 two albinic males of Chlosyne nycteis (Dbldy.) were collected along 
a woodland trail on water company land near Lake Gaillard, North Branford, New 
Haven Co., Connecticut. One specimen has the usual orange-red ground color entirely 
replaced by white (Fig. 1). The other is only partially albinic, with creamy overlay in 
the hindwing medial and forewing subapical areas, and on wing bases. In both, the 
melanic border and markings are apparently unaltered. These specimens were taken 
in the company of typical orange-red C. nycteis. Some 10 to 15 adults were seen in 
the area on that day. A third specimen, also fully albinic, was collected 1 July 1979 at 
the same locality by William Martha. We are aware of no other wild-captured albinic 
C. nycteis. 

The white color may represent the expression of an extremely rare allele akin to 
“whitish” and “blonde” of Colias (see Remington, 1954, Lepid. News 7:139-145). That 
the specimens were collected from within a small, local population of C. nycteis also 
suggests they may be the progeny of a single female. Oliver (1979, J. Lepid. Soc. 32: 
309) and Shapiro (1966, Butterflies of the Delaware Valley, Spec. Publ. Amer. Entomol. 
Soc. 20) have reported albinos of Phyciodes tharos Drury from Pennsylvania. It is of 
interest to note the occurence of similar white phenotypes in these two very closely 
related butterfly genera. Albinics have yet to segregate out of mass cultures of C. 
nycteis, C. harrisii (Scudder), and Phyciodes spp. (C. G. Oliver, pers. comm.). Our two 


86 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Fic. 1. Albinic variants of Chlosyne nycteis. A, Dorsal surfaces. From top: normal 
male, partial albinic, total albinic; B, Ventral surfaces of same specimens as in A. 
Specimens collected 8 July 1979, near Lake Gaillard, North Branford, New Haven Co., 
CT, leg. L. F. Gall & D. F. Schweitzer. Color filter used to enhance contrast between 
albinics and normal male. 


specimens have been deposited in the entomological collections at the Peabody Mu- 
seum of Natural History, Yale University. 


LAWRENCE F.. GALL AND DALE F. SCHWEITZER, Department of Biology, and Ento- 
mology Section, Peabody Museum, Yale University, New Haven, Connecticut 06520. 


VOLUME 37, NUMBER 1 87 


Journal of the Lepidopterists’ Society 
37(1), 1983, 87-88 


THYRIS MACULATA (THYRIDIDAE) AND THREE SPECIES OF 
CLEARWING MOTHS (SESIIDAE) ATTRACTED TO 
AN ARTIFICIAL CHEMICAL BAIT 


Attraction of female sex pheromones for males of many members of the Sesiidae is 
well known (Sharp et al., 1978, Florida Entomol. 61(4):245-250; 1979, Entomol. Soc. 
Amer. Symposium “Pheromones of Sesiidae.”” ARR-NE-6:35-46; Reed et al., 1981, 
Environ. Entomol. 10(4):488-491), and it was interesting to discover that a moth rep- 
resenting another family is also attracted to at least one of these compounds. I spent 
most of July and August of 1980 collecting male clearwing moths with the aid of a sex 
attractant containing mostly the Z,Z isomer of 3,13-octadecadien-l-ol acetate (Z,Z- 
ODDA), a main component in the pheromone system of many sesiids. For a review of 
a sex attractant study see Duckworth and Ejichlin (1977, J. Lepid. Soc. 31:191-196). 
The lure was kindly provided for my use by John Holoyda, whose interest in the 
Sesiidae induced me to participate in this study. The lure strip was enclosed in a piece 
of nylon netting material and pinned to the frame of my collecting net. 

The study sites were mostly virgin prairie remnants in northeastern Illinois that have 
miraculously escaped the plow and the bulldozer, which years ago converted most of 
the Grand Prairie of Illinois to man’s commercial uses. On 6 July I entered Harlem 
Hills Nature Preserve, a 53-acre dry upland prairie in Winnebago Co. Situated on the 
outskirts of Rockford, the prairie was dedicated as a Nature Preserve in 1973. Although 
surrounded by residential development it remains a relatively intact pathway into II- 
linois’ past, when the State abounded with grasses and wildflowers of every descrip- 
tion. About 0930 to 0940 CDT I observed several sesiids hovering around my net and 
zeroing in on the attractant. I netted specimens of two species, subsequently identified 
as Carmenta anthracipennis (Bdv.) and Albuna fraxini (Hy. Edw.), the former being 
the more common of the two. Individuals were so intent on locating the source of the 
lure that they paid scant attention to my efforts with the net. 

While attending to some freshly captured specimens, I observed a very small moth 
near my net, hovering in much the same manner as the two sesiid species just en- 
countered. After netting and examining the tiny specimen, I saw that it was not a sesiid 
but rather a moth of a different family. It was latter identified as Thyris maculata 
(Harris). Though apparently wide ranging from eastern United States through Texas 
into Mexico, this thyridid species is uncommon in collections (from unpublished data). 

Before leaving the prairie site I collected one additional specimen of T. maculata, 
a few more A. fraxini specimens, and a long series of C. anthracipennis. 

The day was intermittently sunny and overcast, but the changes in cloud cover did 
not seem to affect the activity of the male sesiids coming to the bait. Without the use 
of the attractant this would not have been the case, I’m certain; I have never observed 
clearwing moths nectaring on any but bright sunny days. Later on in the summer I 
collected specimens of C. anthracipennis in Lake, Cook and McHenry Counties in 
northeastern Illinois, all from similar prairie habitats and all captured when attracted 
to the same lure. This sex attractant also contributed to the capture of a third sesiid 
species, Synthandon exitiosa (Say), the species from which the Z,Z isomer had been 
extracted, identified and ultimately synthesized. Two specimens of the latter were 
collected on the three acre Cary Prairie, Cary, McHenry Co. on 7 August. This tiny 
parcel of natural prairie is situated in the middle of town, surrounded by homes and a 
Junior High School. 

Specimens of S. exitiosa were also collected at Warren Dunes State Park, Berrien 
Co., Michigan on 26 August. These moths were encountered along the Lake Michigan 
bathing beach, having flown from some place farther inland. 

One specimen of T. maculata has been placed in the collection of the Hlinois Natural 
History Survey (INHS), Urbana, and the other has been retained in my own collection 


88 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


temporarily. Specimens of all three sesiid species have been placed in the INHS col- 
lection, in the collection of John Holoyda, Chicago, and the remainder I have retained. 

It is a pleasure to acknowledge with warm thanks the help given me by Dr. Thomas 
D. Eichlin, Insect Taxonomy Laboratory, Department of Food and Agriculture, Sac- 
ramento, California, in examining and determining the identity of T. maculata and the 
three sesiid species. 


IRWIN LEEUW, 1219 Crystal Lake Road, Cary, Illinois 60013. 


Journal of the Lepidopterists’ Society 
37(1), 1983, 88-89 


EUCOSMOMORPHA ALBERSANA (HUBNER), A PALAEARCTIC SPECIES, 
COLLECTED IN NORTH AMERICA (TORTRICIDAE, GRAPHOLITINI) 


Among undetermined olethreutine moth specimens in the Michigan State University 
Entomology Museum, I discovered a single male of Eucosmomorpha albersana (Hub- 
ner) (Figs. 1, 2). Label data include Midland Co., Mich., June 2, 1961, R. R. Dreisbach, 
genit prep PJ 163. The genus Eucosmomorpha has not previously been reported in 
North America. 

Eucosmomorpha Obraztsov, 1951 is monobasic (Obraztsov, N. S., 1961, Tijd. Ento- 
mol. 104:51-70). Its structural distinctness makes it unlikely to be confused with any 
other genus. The Palaearctic distribution of the one described species. E. albersana, 
is extensive: from the United Kingdom and Scandinavia east into Asia (Bradley, J. D., 
W. G. Tremewan & A. Smith, 1979, British Tortricoid Moths, Tortricidae: Olethreuti- 
nae, London, 336 pp.; Benander, P., 1950, Svensk Insektfauna 10, Tortricina, 173 pp.; 
Bentinck, G. A., Graaf & A. Diakonoff, 1968, Monogr. Nederl. Entomol. Ver. 3, 201 
pp.; Hannemann, H. J., 1961, Die Tierwelt Deutschlands ... 48... Tortricidae, 236 
pp.; Kuznetsov, V. I., 1978, Taxonomic Key to Insects of the European USSR, 4, Lep- 
idoptera, 21, Tortricidae, pp. 193-680 (Russian)). 

The Michigan specimen has the forewing more intricately patterned than western 
European examples. It might be E. albersana ussuriana (Caradja, A., 1916, Deut. Ento- 
mol. Z. “Iris” 30:1-88), but no authentic representatives of this taxon were available 
to me for comparison. With a forewing length of 5.5 mm, the specimen is slightly 


Fics. 1,2. Michigan specimen of Eucosmomorpha albersana: 1, forewing pattern; 
2, male genitalia. 


VOLUME 37, NUMBER 1 89 


smaller than Palaearctic examples, whose forewings are usually stated to range from 
6.5 to 7.5 mm long. The male genitalia (Fig. 2) are indistinguishable from those of an 
example in the National Museum of Natural History collected in Kent, U.K. 

Apparently, the Michigan specimen was captured flying or at a light, so its larval 
host plant is unknown. In Europe the larva feeds within tied leaves of Lonicera and 
Symphoricarpos, genera of the Caprifoliaceae or honeysuckle family (Swatschek, B., 
1958, Die Larvalsystematik der Wickler, Berlin, 269 pp.). 

Whether the specimen represents a population now extant or extinct, introduced or 
endemic, is thus far undetermined. There has been no confirmation in two decades, 
and, although far from traditional ports of entry, the collection area is near Great Lakes 
routes of international shipping through the St. Lawrence Seaway. 


WILLIAM E.. MILLER, North Central Forest Experiment Station, 1992 Folwell Ave- 
nue, St. Paul, Minnesota 55108. 


Journal of the Lepidopterists’ Society 
37(1), 1983, 89 


MELIPOTES INDOMITA (WALKER) IN HAWAII 


In this Journal, vol. 33(2):136, was a note concerning this species in Hawaii, which 
very easily could be understood as if it were a first report of it in the Islands. This, 
however, is not so. Melipotes indomita was reported for the first time on 8 June 1969 
on a building wall in Manoa and then repeatedly at Hickam Air Force Base and Ho- 
nolulu Airport. By 7 August the moth had been found already on Molokai and, shortly 
before that date on Kauai, and in September also on Maui. Now the moth is one of the 
most common noctuids.in the Islands, which is understandable because of the abun- 
dance of the foodplant, the monkeypod tree (Samanea saman (Jacq.)). A very thorough 
description and the life cycle of the moth was published by Oda & Mau (1972, Proc. 
Hawaiian Entomol. Soc. 21(3):435-441). 


J. C. E. RIoTTE, Research Associate, Entomology, B. P. Bishop Museum, P.O. Box 
19000-A, Honolulu, Hawaii 96819. 


Journal of the Lepidopterists’ Society 
37(1), 1983, 89-90 


TMOLUS AZIA (LYCAENIDAE) AND ANTEOS CHLORINDE (PIERIDAE) 
IN THE DOMINICAN REPUBLIC 


Tmolus azia (Hewitson) has recently been collected in Jamaica, the first record of 
its occurrence in the Caribbean (Vyhmeister, G., 1980, J. Lepid. Soc. 34(1):60). On 22 
June 1981 at least three members of the Lepidopterists’ Society Dominican Republic 
expedition collected single specimens of this butterfly in the “desert” region of San- 
tiago Province, approximately 10 km NW of the city of Santiago and several hundred 
meters from the north bank of the Rio Yaque del Norte. This collection date followed 
approximately 40 days of rain, and the local vegetation was lush and dense. The col- 
lectors were Andrew F. Beck, S. S. Nicolay and Charles Zeiger; S. S. Nicolay identified 
the specimens. Mr. Nicolay returned to this site on 28 June 1981 and collected three 


90 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


additional specimens, suggesting strongly that this insect is established at this locale 
and is not merely a visitor. : 

In addition, I collected a single, fresh male Anteos chlorinde (Godart) in Jarabacoa, 
La Vega Province, in the afternoon of 24 June 1981. It was captured on Hibiscus 
blossoms along a roadside in the vicinity of the Hotel Pinar Dorado. Riley (1975, A 
Field Guide to the Butterflies of the West Indies, Demeter Press) indicates that this 
butterfly is not recorded from Hispaniola. 


ANDREW F. BECK, Department of Entomology, Virginia Polytechnic Institute and 
State University, Blacksburg, Virginia 24061. 


Journal of the Lepidopterists’ Society 
37(1), 1983, 90 


HAND-PAIRING OF BATTUS PHILENOR (PAPILIONIDAE) 


Hand-pairing is a useful technique in the laboratory rearing of butterflies (Clarke & 
Sheppard, 1956, Lepid. News 10:47-53) and has been applied in several families, most 
extensively in the Papilionidae. I have successfully used it for most eastern North 
American swallowtails but had been unable to hand-pair Battus philenor (L.) Recent 
observations of mating of caged B. philenor revealed the probable reason for my failure 
and suggested how I might be able to hand-pair the species, which has now been 
accomplished. 

B. philenor males and females were released within a day of eclosion in a large 
outdoor flight cage (7.6 m x 4.6 m x 4.6 m high). Within a few minutes the following 
behavioral sequence was observed for two pairs: A motionless female sitting upright 
on a honeysuckle stem about 3 m from the ground was approached by a male, which 
quickly landed upside down on the stem beneath the female and in a few seconds had 
curved his abdomen up and locked in copula. He immediately hung free with folded 
wings in the usual way. Two aspects of this sequence were unexpected: There was no 
fluttering of wings by either individual; and the male approached from below with his 
body initially parallel to that of the female in a frontal position. 

The usual technique in hand-pairing is to bring the tips of the abdomens together 
at about 180°, squeeze the male to open his claspers and join the two. If this technique 
is tried with B. philenor, both individuals curve the tips of their abdomens under, and 
the two cannot be joined. The observations in the flight cage suggested that the cur- 
vature facilitates copulation in the natural position. I, therefore, brought together a 
male and female in the frontal position, pressed the male’s abdomen to open his clasp- 
ers, and the two immediately joined. Several pairs were so mated and dissection re- 
vealed a spermatophore in each female. B. philenor seems to remain in copula some- 
what longer than most other swallowtails, but, as in other species (Clarke & Sheppard, 
1956, op. cit.), about 20 minutes is sufficient for the passage of a spermatophore. 


Davip A. WEsT, Department of Biology, Virginia Polytechnic Institute and State 
University, Blacksburg, Virginia 24061. 


VOLUME 37, NUMBER 1 9] 


Journal of the Lepidopterists’ Society 
37(1), 1983, 91 


SPECIES OF EUCALYPTUS AS FOOD PLANTS 
FOR LEPIDOPTERA IN EAST AFRICA 


Dr. I. F. B. Common’s paper, “Some factors responsible for imbalances in the Aus- 
tralian fauna of Lepidoptera’ (1981, J. Lepid. Soc. 34:286—294), and his remarks on the 
role of Eucalyptus as a lepidopterous food plant suggest that a list of the Lepidoptera 
feeding on the introduced Eucalyptus spp. in East Africa might be of interest. 

Various species of Eucalyptus are grown in many parts of East Africa, primarily as 
sources of firewood but also as useful agents in the reclamation of swampy land. E. 
citriodora is grown as a plantation crop in Zaire and elsewhere for its oil, used in the 
perfumery trade. 

The following is a list of species recorded as feeding on species of Eucalyptus: 
Lymantriidae—Euproctis molunduana Auriv., Dasychira georgiana Fawcett, D. basa- 
lis Wlk., Argyrostagma niobe Weymer; Lasiocampidae—Lechriolepis nigrivenis Strand, 
Bombycopsis bipars Wlk., Nadiasa cuneata Distant, Pachypasa subfascia Wlk., P. pa- 
pyri Tams, Eucraera salambo Vuillot; Saturniidae—Bunaea alcinoe Stoll, Nudaurelia 
conradsi Rebel, N. cytherea F., N. dione F., N. krucki Hering, N. gueinzii Karsch, 
Lobobunaea phaedusa Drury, Urota sinope Westwood, Athletes ethra Westwood; No- 
todontidae—Desmeocraera varia Janse; Limacodidae—Latoia chapmanni Kirby; 
Psychidae—Eumeta rougeoti Bourgogne, Kotochalia junodi Hylaerts; Noctuidae— 
Euxoa longidentifera Hampson, Spodoptera littoralis Bdv., Heliothis armigera (Hbn.), 
Anua mejanesi Gn., A. tirhaca Cramer, Achaea lienardi Bdv., A. catella Gn., A. faber 
Holland, Plusia limbirena Gn.; Geometridae—Orthonama obstipata F., Colocleora 
divisaria Wlk., Ascotis selenaria Schiff., A. reciprocaria Wlk., Cleora nigrisparsalis 
Janse, C. herbuloti Fletcher, C. dargei Herbulot, C. scobina Fletcher, C. rothkirchi 
Strand, Luxiaria curvivena Warren; Pyralidae—Herculia tenuis Butler, Sylepta bal- 
teata F. As shown for Australia no African butterfly larva have been known to feed on 
Eucalyptus. 

The two indigenous genera of the Myrtaceae, Eugenia and Syzigium, serve as host 
plants for relatively few lepidopterous larvae: five Charaxes spp., one lycaenid, two 
lymantriids, one each lasiocampid, thaumetopoeid and metarbelid, three limacodids 
and two noctuids; however, another introduced genus, Psidium, originally from tropical 
America, is eaten by two lycaenids, one lymantriid, one lasiocampid, six saturniids, 
three notodontids, one limacodid, one metarbelid, one noctuid and two geometrids. 

It is most unusual for introduced species to be eaten by more species than the 
indigenous plants. In the Mimosaceae, for example, there are no records of lepidop- 
terous larvae feeding on the introduced Leucaena glauca and Acacia decurrens, nor 
are the flowers visited by imagines; yet, the indigenous species of Acacia are eaten by 
numerous species of lepidopterous larvae, and their flowers are highly attractive to 
butterflies and moths. 


D. G. SEVASTOPULO, F.R.E.S., P.O. Box 95617, Mombasa (Nyali), KENYA. 


Journal of the Lepidopterists’ Society 
om), 1983, 91-92 


SUMMER BUTTERFLIES IN DINOSAUR NATIONAL MONUMENT 


The canyons of the Green River in Dinosaur National Monument are in large part 
accessible only by raft. They are in a most interesting area biogeographically, however, 


92 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


lying just off the eastern end of the Uinta Mountains, the most southern outpost of the 
northern Rocky Mountains. To the north and east of Dinosaur lies the Wyoming Basin, 
which separates the northern and southern Rockies and forms a gap that has not been 
crossed by such butterflies as Euphydryas gillettii (Barnes) and Parnassius clodius 
Ménétriés. 

A raft trip down the Green River in mid-July of 1981. provided an opportunity for a 
quick survey of the butterfly fauna of the canyons. Although during the trip (12-17 
July) the river was cold, the desert floor of the canyons was hot, and few wildflowers 
were in bloom. In general the butterfly fauna was unexciting. Asclepias was abundant, 
and Danaus plexippus L. was the most prominent butterfly. Occasional individuals of 
Vanessa cardiu (L.) and Pieris rapae (L.) were seen, and once or twice a day a Papilio 
multicaudatus Kirby would fly by. In dry grassy areas Cercyonis oetus (Bdv.) was 
frequent, second only to monarchs in abundance. Occasionally an Erynnis afranius 
(Lintner) was seen. A single Colias flew by, possibly alexandra Edw. 

Near Upper Disaster Falls in the Lodore Canyon, Colorado (mile 237) some thistles 
were in bloom. While our party was reconnoitering the rapids I took a couple of Spey- 
eria coronis Behr and then was attracted by the unusual silhouette of a backlighted 
“monarch” nectaring. A second look told me it was no monarch—indeed it was Spey- 
eria nokomis (Edw.), a species I had not seen before. Two males were taken of this 
scarce and strangely distributed fritillary—previously unknown in northwestern Col- 
orado but present in adjacent northeastern Utah (Uintah County—Callaghan and Tid- 
well, 1972, J. Res. Lepid. 10:191—202; Ferris & Fisher, 1971, J. Lepid. Soc. 25:44—52). 

Since mineral development is rampant in Uintah County, it is a pleasure to report 
this minor range extension, because it means that S. nokomis is established in a national 
monument where its habitat will be protected. It is clear that further exploration of the 
canyons of Dinosaur National Monument would be useful, to establish both the nature 
of the butterfly fauna earlier in the season and the extent of the S. nokomis colony. 

L. P. Grey kindly identified the Speyeria coronis; John M. Burns the Erynnis. 


PAUL R. EHRLICH, Department of Biological Sciences, Stanford University, Stan- 
ford, California 94305. 


Journal of the Lepidopterists’ Society 
37(1), 1983, 92-94 


NEW BUTTERFLY DISTRIBUTION RECORDS FOR 
NORTHERN NEW YORK STATE 


In general, the Adirondack and St. Lawrence valley regions of New York are poorly 
represented in entomological collections owing, in part, to a dearth of resident collec- 
tors in these regions. This is especially evident in the lack of records of many species 
of Lepidoptera from these areas. The following annotated list of Rhopalocera supple- 
ments Shapiro’s list of the butterflies and skippers of New York (1974, Search: Agri- 
culture, Cornell Univ. Agric. Exp. Sta. No. 12, Ithaca, N.Y., 60 pp.). 

Most specimens were collected or observed at White’s Hill (elev. 438 m), Parishville, 
in St. Lawrence County. White’s Hill lies just within the northwestern boundary of the 
Adirondack Park and overlooks the St. Lawrence valley to the north. This area is, 
therefore, intermediate between the lower, very flat plain extending to the St. Law- 
rence River and the much higher Adirondack Mountains. Some areas of White’s Hill 
are old fields in succession, but much of the land is second-growth deciduous forest, 
with pockets of hemlock and spruce. 

Additional records within St. Lawrence Co. are presented from the Cranberry Lake 
Biological Station (CLBS), owned by the State University of New York College of 
Environmental Science and Forestry; and Sterling Pond, located about 13 km SE of 


VOLUME 37, NUMBER 1 93 


White’s Hill. Other records are given for Onondaga Co. and Dutchess Co. New county 
records are designated by an asterisk (*). The format follows Shapiro (1974, op. cit.). 
The specimens are in the author's collection. 

Satyridae 

Lethe anthedon (Clark)—St. Lawrence Co.*, Parishville, 12-28 July. In 1974-75, this 
species was semi-abundant in rocky, sunlit deciduous woods. 

Lethe eurydice (Johansson)—Dutchess Co.*, Amenia, July 1977. St. Lawrence Co., 
Parishville, 8-20 July. Very common locally in wet meadows, sighted frequently at 
CLBS in old beaver meadows. 

Coenonympha tullia inornata Edwards—St. Lawrence Co.*, Parishville, 23 June 
1973; 11-17 June 1975; 23-24 Aug. 1973, 75. Most often encountered in wet meadows, 
and two broods per year are apparent. 

Euptychia cymela (Cramer)—St. Lawrence Co.*, Parishville, Sterling Pond, and CLBS, 
9-25 June. Collected along margins of open deciduous woods. 

Cercyonis pegala nephele (Kirby)—St. Lawrence Co.*, Parishville, 26 July 1974. In 
dry sloping, abandoned field. Only occasional specimens have been seen in the area. 
This specimen does not appear to represent an intergrade between C. pegala alope 
Fabr. and C. pegala nephele. 


Nymphalidae 


Speyeria cybele (Fabricius)—This common and widespread species has been col- 
lected frequently at Parishville and CLBS during July. 

Speyeria atlantis (Edwards)—St. Lawrence Co., CLBS, 26 July—15 Aug. Not a par- 
ticularly common species. 

Speyeria aphrodite (Fabricius)—St. Lawrence Co., Parishville, August. Common in 
dry, brushy fields feeding at composites. 

Boloria selene myrina (Cramer)—St. Lawrence Co., Parishville, 22 June 1974. In 
open marshy area with exposed patches of flat rock surrounded by small spruces and 
red maples. 

Euphydryas phaeton (Drury)—St. Lawrence Co., Parishville, late June to mid-July. 
Not particularly abundant, even where its foodplant (Chelone glabra L.) is found. 

Chlosyne harrisii (Scudder)—St. Lawrence Co., Parishville, 11-22 June, in boggy 
meadows. Collected further northward than previously recorded in New York. 

Polygonia interrogationis (Fabricius)—St. Lawrence Co.*, Parishville, 9 Sept.; CLBS, 
2 Aug. Onondaga Co., Lafayette Experiment Station, 20 June. The St. Lawrence Co. 
records extend the range more northward than previously recorded for the state. It 
seems doubtful that the adults overwinter in St. Lawrence Co. 

Polygonia comma (Harris)—St. Lawrence Co.*, Parishville, 23 May 1975. In open 
woods near a long-abandoned farm. Wild hops (Humulus sp.) growing nearby may 
have been the foodplant. 

Polygonia faunus (Edwards)—St. Lawrence Co., Parishville, 7 Sept. Collected along 
wooded margins of a dirt road. Other records are mostly from the central Adirondacks 
(Shapiro, 1974 op. cit.). 

Nymphalis j-album (Boisduval & LeConte)—St. Lawrence Co., Parishville. The 17 
April is earliest, 25 Oct., the latest date of collection. 

Nymphalis milberti (Godart)—In Aug. 1977, I saw dozens of individuals flying about, 
sunning themselves, and feeding at flowers and spots of moisture on the summit of 
Algonquin Peak (elev. 1558 m) in Essex Co. Shapiro (1974, op. cit.) described this as a 
lowland species. 

Nymphalis antiopa (Linnaeus)—In Syracuse (Onondaga Co.), larvae have caused 
mild defoliation of Hackberry trees (Celtis occidentalis L.) in ornamental plantings. 
Earliest sighting was 23 April in Parishville. 

Vanessa atalanta (Linnaeus)—Not seen during spring months at Parishville, but only 
from mid- to late summer. 

Vanessa cardui (Linnaeus)—St. Lawrence Co., Parishville, 24 May-6 June. I have 
seen this species only three times at White’s Hill. The specimen captured on May 24 
was feeding at Dandelion flowers (Taraxacum officinale Weber). 


94 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Vanessa virginiensis (Drury)—St. Lawrence Co., Parishville. Locally abundant in 
open meadows, from mid- to late summer. 

Asterocampa celtis (Boisduval & LeConte)—Onondaga Co.*, Lafayette Experiment 
Station, and Clark Reservation State Park, 15-30 June 1978, 79. Common along Hack- 
berry-lined woodland roads. Previously sepoured! from a feu southern-most counties 
bordering Pennsylvania and on Long Island (Shapiro, 1974, op. cit.). 

Lycaenidae 

Harkenclenus titus (Fabricius)—St. Lawrence Co.*, Parishville, CLBS, late June to 
early July. Commonly collected at Asclepias sp. flowers in open brushy fields. 

Satyrium liparops (Boisduval & LeConte)—St. Lawrence Co., Parishville, 19 July. 
Collected along fencerow in an old field. 

Callophrys niphon (Hubner)—St. Lawrence Co.*, Parishville, 18-24 May; Oneida 
Co., Boonville, 25 May. Pinus strobus L. was the only pine in the areas where speci- 
mens were taken. Shapiro (1974, op. cit.) suspected that C. niphon may utilize white 
pine where hard pines (P. rigida Mill.) do not occur. 

Lycaena thoe (Guérin-Méneville)—St. Lawrence Co., Parishville, CLBS, mid-July. 
These captures confirm its presence in the western Adirondacks. 

Celastrina argiolus pseudargiolus (Boisduval & LeConte)—Very common in May at 
Parishville, especially along margins of dirt roads. May 11 is the earliest date of capture 
for dd and @Q. 

Papilionidae 

Papilio polyxenes asterias Stoll—St. Lawrence Co., Parishville, 15 Aug. Not com- 
monly seen in this area. There are few records from the northern-most counties. 

Papilio glaucus Linnaeus—The small canadensis form is found in St. Lawrence Co.*, 
Parishville, and CLBS, with May 22 as the earliest capture date. On June 12, 1978, I 


observed large numbers (over 20/min) at White’s Hill. Many individuals were feeding 
at Orange Hawkweed blossoms (Hieracium aurantiacum L.). 


Pieridae : 

Pieris napi oleracea Harris—Onondaga Co.*, Manlius, 3 May 1977, in open decid- 
uous woods. St. Lawrence Co., Parishville, 17 May—27 June. Commonly collected along 
wooded roads. 

Colias eurytheme Boisduval—St. Lawrence Co., Parishville, 17 Aug.—26 Sept. Not 


very common around the Parishville area (1973-78), and there are few reported local- 
ities for northern New York. 
Hesperiidae 

Erynnis icelus (Scudder & Burgess)—St. Lawrence Co., Parishville, 20 May—21 June. 
Common along wooded dirt roads. It is doubtful that E. brizo Boisduval & LeConte 
occurs in this area, as there are no oaks upon which to feed. 

Erynnis baptisiae (Forbes)—Onondaga Co.*, Syracuse, 20 May. 

Carterocephalus palaemon mesapano (Scudder)—St. Lawrence Co., CLBS, 4 June, 
in a grassy beaver meadow near a spruce bog. 

Ancyloxipha numitor (Fabricius)—St. Lawrence Co.*, CLBS, July. In wet grassy 
areas. 

Polites mystic (Scudder)—St. Lawrence Co., Parishville, 18 July. In a wet meadow. 

Poanes hobomok (Harris)—St. Lawrence Co., Parishville, CLBS, 26 May—18 June. A 
very common skipper in the county. 

Amblyscirtes hegon (Scudder)—St. Lawrence Co.*, Parishville, 29 May. 


ACKNOWLEDGMENTS 


I would like to thank W. H. Wagner, Department of Botany, University of Michigan, 
for checking several identifications, and also I. J. Cantrall, Museum of Zoology, for 
providing helpful comments on the manuscript and offering encouragement. 


MARK F. O'BRIEN, Insect Division, Museum of Zoology, University of Michigan, 
Ann Arbor, Michigan 48109. 


Journal of the Lepidopterists’ Society 
37(1), 1983, 95-96 


BOOK REVIEWS 


THE BUTTERFLIES OF THE MALAY PENINSULA, by A. Steven Corbet and H. M. Pen- 
dlebury, 3rd ed., revised by J. N. Eliot. 1978. Malayan Natural History Society, Kuala 
Lampur, Malaysia. v—xiv + 578 pp., ill. 


Col. Eliot has done a magnificant job of revising “the” standard reference to Malay- 
sian butterflies, which was no easy task, inasmuch as the original Corbet and Pendle- 
bury treatment produced perhaps the finest regional text available anywhere. Eliot has 
thoroughly modernized the treatments of all groups of butterflies, save most of the 
Hesperiidae which are left nearly as Evans had classified them in 1949. Among the 
new taxa proposed by Eliot in this book (a practice that I do not like, because such 
descriptions are better made available in journals) are five hesperiid genera, and since 
most new taxa add to clarity, the book is helped by them. 

Introductory treatments of the morphology of various stages (could have been a bit 
more detailed), nomenclature and classification, geographical distribution of butterflies, 
wing patterns, duplex species, sex ratios and the history of collecting on the peninsula 
lead off the book. All sections are well done and the information presented is sound. 

The discussions of species and higher categories are competent and comprehensive. 
As is the case with so many modern books, the nomenclature is not so conservative as 
that in the earlier literature; but in this case, the reasons behind these nomenclatorial 
changes are well elucidated. No reviewer can be expected to wholeheartedly agree 
with everything in a book; this book and this reviewer are not exceptions. On page 
128, Eliot mentions my 1968 higher classification of the Satyridae, stating that it “... 
ignores the male genitalia, takes little account of secondary sexual characters and is 
not adopted here.” Male genitalia were not considered in the revision chiefly because 
the genitalia of Satyridae are so simplified that they offer very few good characters for 
higher classification; for generic and specific determination, they are very appropriate. 
The use of male secondary sexual characters, so abused by Moore and others around 
the tum of the century as the basis of new genera, were not used mainly because 
geneticists tell us that a characteristic can be shifted in position on a wing through the 
action of a single gene, hardly enough to warrant a change in tribal status, the lowest 
point to which my classification went. 

The larvae of some Neotropical Morphidae feed on monocots (others on dicots), 
which characteristic makes that family a pivotal one in the evolution of the nymphalid 
complex. At this time, though, I am inclined to consider at least part of the Morphidae 
(Taenaris, etc.) to perhaps be satyrids, and were I doing the revision today, I should 
consider the genus Penthema to be a satyrid in the Elymniinae, as suggested by Jap- 
anese correspondents. 

I am pleased to see that Cethosia is unequivocally placed in the otherwise Neotrop- 
ical Heliconiinae. This genus sits in the Heliconiinae in about the same position (that 
of an outlier of a Neotropical group) as does Tellervo in the Ithomiidae. 

Lycaenid classification has benefitted in this book by Eliot’s use of his own superb 
higher classification of that family. Classification of members of that family has always 
been a problem with traditional treatments, and the original Corbet and Pendlebury 
handling of the group was no exception. The success Eliot has had with his classifi- 
cation should send a signal to the rest of us that it works, better than any other. It 
should be expected to treat the Malaysian fauna well (it was based chiefly on Malaysian 
examples), but it is certainly adaptable to lycaenids from other regions. 

A brief word about other features of the book must include mention of the very useful 
keys scattered throughout the text. Many amateurs do not like keys, preferring to rely 
instead on illustrations, preferably colored, but keys are the most economical way of 
presenting differences (and similarities) between taxa. These keys are much expanded 
over the original Corbet-Pendlebury ones, thus adding to the utility of the volume for 
both professional and amateur alike. 

For those who admittedly refuse to use keys, there are high quality illustrations as 
well. Not all of the species are figured (nor are their genitalia in the plates of those), 


96 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


but the illustrations are of good quality and clarity. In conjunction with the keys, the 
reader should be able to identify any Malaysian butterfly from the species illustrated 
on the plates. 

In short, this book should be on the shelves of anyone with an interest in the but- 
terflies of this region, even if he has another edition of Corbet and Pendlebury. Eliot 
has done a fine revision that would make both Corbet and Pendlebury happy to ‘have 
their names associated with, and nothing finer can be said about a revision of an older 


book. 


LEE D. MILLER, Allyn Museum of Entomology, 3701 Bay Shore Road, Sarasota, 
Florida 33580. 


Journal of the Lepidopterists’ Society 
37(1), 1983, 96 


THE CUTWORM MOTHS OF ONTARIO AND QUEBEC by Eric W. Rockburne and J. Donald 
Lafontaine, with photographs by Thomas H. Stovell. 1976. Agriculture Canada, Re- 
search Branch, Publication 1593. 164 pp., 613 col. figs. Obtainable from Canadian 
Government Publication Centre, Supply and Services Canada, Hull, Quebec K1A OS9. 
$US 10.50, $CAN 8.50. 


This hard-bound, lavishly illustrated little book is primarily a collection of life-size 
colored illustrations of noctuid moths from the Agaristinae to the Catocalinae (in the 
sequence of the McDunnough, 1938, check list), intended as an identification guide 
for the amateur. It has a brief text of usually three or four lines per species, giving 
distribution within the two provinces, food plants, and flight period, and a 5-page 
introduction consisting of elementary information on classification, life history, adult 
structure, and collecting. It treats all Noctuidae represented by specimens from Ontario 
or Quebec in the Canadian National Collection with the exception of the Hypeninae, 
Rivulinae, and Herminiinae (in the sense of McDunnough). My only complaint about 
the book is that these latter three also could have been covered with the addition of 
only two and one-half pages to the 41 pages of illustrations already included, making 
it a nearly complete guide to the Noctuidae of that region. The term “cutworm moths” 
is construed.as encompassing such a large part of the Noctuidae that it might just as 
well have been applied to them all. 

This book is essentially without errors, and the nomenclature is current as of the 
time of publication. The colored illustrations are not perfect, but they would have to 
be regarded as adequate to excellent when one considers the modest price. The original 
photographs, made against the traditional pale-blue background by the same photog- 
rapher who did the illustrations for the Butterflies and Moths of Newfoundland and 
Labrador, were obviously very well done. As in the work just cited, the legends give 
no locality data, but the stated purpose of Rockburne and Lafontaine was only to pro- 
duce “a handbook intended for amateurs.” Also, I could see nothing in the illustrations 
to reveal that the figured specimens were from anywhere other than Ontario or Quebec. 

I do not hesitate to recommend this book as a useful aid to the identification of 
noctuid moths of the Northeast and suspect that it will find an important place on the 
bookshelves of many entomologists who do not think of themselves as amateurs. 


DOUGLAS C. FERGUSON, Systematic Entomology Laboratory, IIBII, Agricultural 
Research Service, U.S.D.A., % U.S. National Museum of Natural History, Washington, 
D.C. 20560. 


Date of Issue (Vol. 37, No. 1): 19 August 1983 


EDITORIAL STAFF OF THE JOURNAL 
THOMAS D. EICHLIN, Editor 


% Insect Taxonomy Laboratory 
1220 N Street 
Sacramento, California 95814 U.S.A. 


MAcDA R. Papp, Editorial Assistant 


_ DoucLas C. FERGUSON, Associate Editor THEODORE D. SARGENT, Associate Editor 


: NOTICE TO CONTRIBUTORS 


_ Contributions to the Journal may deal with any aspect of the collection and study of 
Hes “Lepidoptera Contributors should prepare manuscripts according to the following instruc- 
_ tions. 
Abstract: A brief abstract should precede the text of all articles. 
Text: Manuscripts should be submitted in triplicate, and must be typewritten, en- 
 tirely double-spaced, employing wide margins, on one side only of white, 8% X 11 inch 
Bisaiver. Titles should be explicit and descriptive of the article’s content, including the 
_ family name of the subject, but must be kept as short as possible. The first mention of a 
_ plant or animal in the text should include the full scientific name, with authors of 
R zoological names. Insect measurements should be given in metric units; times should be 
| given in terms of the 24-hour clock (e.g. 0930, not 9:30 AM). Underline only where 
_ italics are intended. References to footnotes should be numbered consecutively, and the 
footnotes typed on a separate sheet. 
Literature Cited: References in the text of articles should be given as, Sheppard 
(1959) or (Sheppard 1959, 1961a, 1961b) and all must be listed alphabetically under the 
heading LITERATURE CITED, in the following format: 


SHEPPARD, P. M. 1959. Natural selection and heredity. 2nd. ed. Hutchinson, London. 
209 pp. 

196la. Some contributions to population genetics resulting from the study of 

the Lepidoptera. Adv. Genet. 10: 165-216. 


In the case of general notes, references should be given in the text as, Sheppard (1961, 
Adv. Genet. 10: 165-216) or (Sheppard 1961, Sym. R. Entomol. Soc. London 1: 23-30). 
Illustrations: All photographs and drawings should be mounted on stiff, white back- 
ing, arranged in the desired format, allowing (with particular regard to lettering) for 
reduction to their final width (usually 4% inches). Illustrations larger than 8% X 11 inches 
are not acceptable and should be reduced photographically to that size or smaller. The 
author's name, figure numbers as cited in the text, and an indication of the article's title 
should be printed on the back of each mounted plate. Figures, both line drawings and 
halftones (photographs), should be numbered consecutively in Arabic numerals. The term 
“plate” should not be employed. Figure legends must be typewritten, double-spaced, on 
a separate sheet (not attached to the illustrations), headed EXPLANATION OF FIGURES, 
with a separate paragraph devoted to each page of illustrations. 
Tables: Tables should be numbered consecutively in Arabic numerals. Headings for 
tables should not be capitalized. Tabular material should be kept to a minimum and 
must be typed on separate sheets, and placed following the main text, with the approx- 
imate desired position indicated in the text. Vertical rules should be avoided. 
Proofs: The edited manuscript and galley proofs will be mailed to the author for 
correction of printer’s errors. Excessive author’s changes at this time will be charged to 
authors at the rate of 75¢ per line. A purchase order for reprints will accompany the 
proofs. 
Correspondence: Address all matters relating to the Journal to the editor. Short 
manuscripts such as new state records, current events, and notices should be sent to the 
editor of the News: June Preston, 832 Sunset Drive, Lawrence, Kansas 66044 U.S.A. 


PRINTED BY THE ALLEN PRESS, INC., LAWRENCE, KANSAS 66044 U.S.A. 


CONTENTS 


TERRITORIAL BEHAVIOR OF NYMPHALIS ANTIOPA AND POLYGONIA 
COMMA (NYMPHALIDAE). Royce J. Bitzer & Kenneth C. 


A NEw SPECIES OF AGROTIS OcHs. (Noctuidae) from Sable Is- 
land, Nova: Scotia: Kenneth Neil 0 


A NEw SPECIES OF SHINIA (NOCTUIDAE) FROM MANITOBA AND 
SASKATCHEWAN WITH DESCRIPTION OF ITS LIFE 
History. D: F. Hardwick 0 


A NEw GENUS AND NEW SPECIES OF GEOMETRID MOTH FROM 
TEXAS. Douglas C.. Ferguson 2. 


PROLONGED DIAPAUSE AND PUPAL SURVIVAL OF PAPILIO ZELI- 
CAON LUCAS (LEPIDOPTERA: PAPILIONIDAE). Steven R. 


NOTES ON THE BIOLOGY OF AGONOPTERIX ALSTROEMERIANA 
(CLERCK), WITH DESCRIPTIONS OF THE IMMATURE STAGES 
(OECOPHORIDAE). M. Berenbaum > S. PASSO -oocccciccccc0cn 


GEOGRAPHIC DISTRIBUTION AND CHECKLIST OF THE BUTTER- 
FLIES OF KERN COUNTY, CALIFORNIA. Ken Davenport .... 


ERBLICHIA ODORATA SEEM. (TURNERACEAE) IS A LARVAL Host 
PLANT OF EUEIDES PROCULA VULGIFORMIS (NYMPHALIDAE: 
HELICONIINI) IN SANTA ROSA NATIONAL PARK, COSTA 


RicA. D. He Janzen 20 70 
GENERAL NOTES 
The suitability of Juniperus (Cupressaceae) for larvae of Callophrys hesseli 
(Rawson and Ziegler) (Lycaenidae). Kurt JOMMSOMN 2... cece 78 
The type of Argynnis apacheana Skinner. F. Martin Brown . 79 
A new food plant record for Chlosyne gorgone carlota (Reakirt) (Nym- 
phalidae).’ David: GC. [finer 2.25 0 a ee 80 
A natural occurrence of inter-tribal copulation in the Papilionidae. Mark 
D. Rausher' d+ M. Berenbaum 20 a ee 81 
Notes on the courtship of Troides oblongomaculatus papuensis (Papilioni- 
dae) in Papua New Guinea. Michael Parsoms nccccccccecececccccvcccnccccsssseeeeeneneeseeee 83 
Albinic variants of Chlosyne nycteis from Connecticut (Nymphali- 
dae): _ Lawrence F. Gall ¢> Dale F. Schweitzer, 02 ee 85 
Thyris maculata (Thyrididae) and three species of clearwing moths (Sesi- 
idae) attracted to an artificial chemical bait. Irwin Leeuw 0000 87 
Eucosmomorpha albersana (Hiibner), a Palaearctic species, collected in North 
America (Tortricidae, Grapholitini). William E. Miller 88 
Melipotes indomita (Walker) in Hawaii. J. C. E. Riotte 2 89 
Tmolus azia (Lycaenidae) and Anteos chlorinde (Pieridae) in the Domini- 
ean Republic. “Andrew F) Beck oo 0s ge 89 
Hand-pairing of Battus philenor (Papilionidae). David A. West 90 
Species of Eucalyptus as food plants for Lepidoptera in East Africa. D. 
G. Sevastapulo: sc Ni PO Ne 2S Us ee nt te 91 
Summer butterflies in Dinosaur National Monument. Paul R. Ehrlich ....... 91 
New butterfly distribution records for northern New York state. Mark F. 
OMBraen ayn NaN NN EIS Og els SO RP 92 
BOOK PREVIEWS) (2 UA Pe NEOUS wey aie a IE Oy BB Ae em 28, 69, 95, 96 
OBITUARY 005) S00 oS Se Ne Sa a ee 37 


a 
Volume 37 1983 Number 2 


vA 


ISSN 0024-0966 


JOURNAL 


_LEPIDOPTERISTS’ SOCIETY 


Published quarterly by THE LEPIDOPTERISTS’ SOCIETY 


Publié par LA SOCIETE DES LEPIDOPTERISTES 
Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN 
Publicado por LA SOCIEDAD DE LOS LEPIDOPTERISTAS 


6 September 1983 


THE LEPIDOPTERISTS’ SOCIETY 


EXECUTIVE COUNCIL 


CHARLES V. COVELL, JR., President LINCOLN P. BROWER, 
CLIFFORD D. FERRIS, Vice President Immediate Past President 
ALBERTO Diaz FRANCES, Vice President JULIAN P. DONAHUE, Secretary 
CLAUDE LEMAIRE, Vice President RONALD LEUSCHNER, Treasurer 


Members at large: 


R. L. LANGSTON K. S. BROWN, JR. F. S. CHEW 
R. M. PYLE T. C. EMMEL _G. J. HARJEs 
A. M. SHAPIRO E. H. METZLER 


The object of the Lepidopterists’ Society, which was formed in May, 1947 and for- — 
mally constituted in December, 1950, is “to promote the science of lepidopterology in ~ 
all its branches, .... to issue a periodical and other publications on Lepidoptera, to facil- ~ 
itate the exchange of specimens and ideas by both the professional worker and the ~ 
amateur in the field; to secure cooperation in all measures’ directed towards these aims. 

Membership in the Society is open to all persons interested in the study of Lepi- 
doptera. All members receive the Journal and the News of the Lepidopterists’ Society. — 
Institutions may subscribe to the Journal but may not become members. Prospective ~ 
members should send to the Treasurer full dues for the current year, together with their — 
full name, address, and special lepidopterological interests. In alternate years a list of 
members of the Society is issued, with addresses and special interests. There are four 
numbers in each volume of the Journal, scheduled for February, May, August and 
November, and six numbers of the News each year. 


Active members—annual dues $18.00 
Student members—annual dues $12.00 
Sustaining members—annual dues $25.00 
Life members—single sum $250.00 
Institutional subscriptions—annual $25.00 


Send remittances, payable to The Lepidopterists’ Society, and address changes to: 
Ronald Leuschner, 1900 John St., Manhattan Beach, California 90266 U.S.A. 


Back issues of the Journal of the Lepidopterists’ Society, the Commemorative Vol- 
ume, and recent issues of the NEWS are available from the Treasurer. The Commem- 
orative Volume, is $6; for back issues, see the NEWS for prices or inquire to Treasurer. 


Order: Mail to Ronald Leuschner, 1900 John St., Manhattan Beach, California 90266 
U.S.A. 


Journal of the Lepidopterists’ Society (ISSN 0024-0966) is published quarterly by the 
Lepidopterists’ Society, a non-profit, scientific organization. The known office of publi- 
cation is 1041 New Hampshire St., Lawrence, Kansas 66044. Second class postage paid 
at Lawrence, Kansas, U.S.A. 66044. 


Cover illustration: Adult of the squash vine borer, Melittia cucurbitae (Harris) (Sesi- 
idae), which occurs in the eastern half of the United States and along the Gulf Coast into 
Vera Cruz, Mexico. The larvae are destructive borers in the vines of various cultivars of 
Cucurbita spp. (squash, pumpkins and gourds). Original drawing by Dr. Charles S. Papp, 
Sierra Graphics & Typography, 1722 J Street #19, Sacramento, CA 95814, USA. 


oun AL. OF 
Toe LeEPIDOPTERISTS’ SOCIETY 


Volume 37 1983 Number 2 


Journal of the Lepidopterists’ Society 
37(2), 1983, 97-105 


NATURAL HISTORY OF SEVEN HAIRSTREAKS IN 
COASTAL NORTH CAROLINA 


SAMUEL M. GIFFORD 
Route 1, Box 606, Manteo, North Carolina 27954 


AND 


PAUL A. OPLER 
Division of Biological Services, U.S. Fish and Wildlife Service, 
Washington, D.C. 20240 


ABSTRACT. Seven species of hairstreaks were observed in their natural hab- 
itats along coastal North Carolina. Also, larvae of each were reared in the laboratory 
from eggs. Various aspects of the butterflies’ natural history are discussed. The species 
studied are: Satyrium calanus falacer (Godart), S. liparops (Leconte), S. kingi (Klots 
& Clench), Calycopis cecrops (Fabricius), Mitoura hesseli Rawson & Ziegler, Incisalia 
henrici (Grote & Robinson), and Fixsenia ontario (Edwards). 


For the last six years (1974—1980), the senior author has observed 
hairstreak behavior in the field on Hatteras Island and Roanoke Is- 
land, Dare County, North Carolina. In addition, larvae were reared 
in captivity from eggs found in nature or obtained from caged females. 
The seven species considered in the present paper are Satyrium ca- 
lanus falacer (Godart), S. liparops (Leconte), S. kingi (Klots & Clench), 
Calycopis cecrops (Fabricius), Mitoura hesseli Rawson & Ziegler, 
Incisalia henrici (Grote and Robinson), and Fixsenia ontario (Ed- 
wards). The junior author has provided comparative notes in the dis- 
cussion of each species based upon prior literature reports. Each 
species is discussed individually and information for each species is 
also presented on Table 1. 


METHODS 


If possible, females were observed in the field in order to gain some 
idea of their host preferences, and in some cases, e.g., I. henrici, eggs 
were collected in the field. In most instances wild caught females 


JOURNAL OF THE.LEPIDOPTERISTS’ SOCIETY 


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VOLUME 37, NUMBER 2 99 


were caged with appropriate host plant material and nectar sources if 
available. Larvae resulting from eggs obtained by either method were 
kept in individual containers and fresh host material was provided 
each day. The date of each molt, pupation and adult eclosion were 
recorded on each container. Larvae were kept on an outside porch 
under natural temperature conditions. Examples of eggs, larvae, and 
pupae of each species were preserved. Reared adults were deposited 
in the Carnegie Museum of Natural History and the Smithsonian In- 
stitution. 


RESULTS 
Satyrium calanus falacer 


Eggs were obtained from females collected on Roanoke Island and 
caged with twigs of bluejack oak (Quercus incana). The egg-bearing 
twigs were inserted into a styrofoam block within a square cage, which 
was hung from a branch in the senior author's yard. They were sep- 
arated to prevent mold. In general, the developmental behavior was 
similar to that of Fixsenia ontario. The falacer eggs hatched a few 
days later than those of F. ontario; concomitantly, the former's host 
(O. incana) flowered and leafed out a few days later than did Quercus 
virginiana. The larvae fed on catkins at first and then switched over 
to young leaves when they appeared. 

The larvae were.dark green and marked with white. The body setae 
were brown. The larvae looked brownish and were accented by the 
white markings. The mature larvae crawled a good deal and ended 
up pupating on the lids or sides of the containers. Pupae were never 
found in the field. Typical developmental times are shown on Table 
1. Adults emerged from 25 May to 4 June. 

S. calanus utilizes only members of the oak (Fagaceae) and walnut 
(Juglandaceae) families as hosts, although Smith (1797) reports haw- 
thorne (Crataegus: Rosaceae) based on a painting by John Abbot. In 
any one area usually only one host genus is utilized. In most situations 
these are various oaks (present paper; Shapiro, 1966, 1974), but hick- 
ory (Carya) or walnuts (Juglans) are utilized in Texas and some por- 
tions of Florida (Kendall, 1964; D. Baggett, pers. comm.). 

It is possible that there are two or more unrecognized sibling species 
currently going under the name calanus, and that these might be host- 
limited (D. Baggett, pers. comm.). 


Satyrium liparops 


On both Hatteras and Roanoke Islands, the larvae of S. liparops 
were the first to hatch from their eggs in the spring. Their host, high 


100 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


blueberry (Vaccinium corymbosum), was also the first shrub in the 
area to put out new spring growth. It may be that the same weather 
conditions govern both events. Eggs and larvae were collected on 
Vaccinium in the field at both localities. Eggs were laid on host twigs 
the previous summer. Female S. liparops lay eggs both on reproduc- 
tive and non-reproductive hosts, but seem to preferentially select por- 
tions of the shrub which will produce the most rapid growth. Eggs 
must be kept under natural moisture conditions outside or they will 
desiccate, and the young larvae will be unable to chew a hole through 
the egg. The remainder of the egg was not eaten. The young larva 
crawled to the nearest bud and bored into it. It fed within the bud 
until the leaves began to unfurl, after which time it fed on the leaves. 
In some instances the larvae bored into flower buds, open flowers and 
developing fruits, all of which were fed upon. The larvae were dark 
green when mature with transverse white markings. The body setae 
were dark brown. Typical developmental times are indicated on Ta- 
ble 1. Adults emerged between 30 May and 6 June. 

This hairstreak is the most catholic Satyrium in its choice of larval 
hosts, although only one or a few may be used in any one region. 
Brown (1976) reported a Florida population to utilize blueberry as its 
host, while Knudsen (1955) reported rhododendron as a host in Geor- 
gia. Thus, southeastern populations may oviposit only on members of 
the heath family (Ericaceae). In contrast, more northern populations 
are known to utilize a wide array of woody hosts, including hawthorne 
(Crataegus), wild plum and cherry (Prunus), shadbush (Amelan- 
chier), blackberry (Rubus), oak (Quercus), willow (Salix), and horn- 
beam (Carpinus) (Klots, 1951; Shapiro, 1974). 


Satyrium kingi 


Found on Roanoke Island, adult females and larvae of S. kingi were 
found in close association or upon sweetleaf or horse-sugar tree (Sym- 
plocos tinctoria: Symplocaceae). This plant occurs in isolated patches 
scattered through the woods on Roanoke Island, where adults were 
also seen nectaring on flowers of chinquapin (Castanea pumila). Eggs 
were placed near twig tips by captive females. The larvae did not eat 
the egg shells after chewing their way out, but bored directly into 
leaf buds and fed there until after the first molt when the leaves began 
to unfurl. Larvae presented with host flowers or fruits refused to feed. 
Two larvae found in nature, when collected with host material for 
captive stock, completed their development at the same time, and the 
resultant adults eclosed on about the same dates. 

The larvae were lighter green than those of S. liparops and were a 


VOLUME 37, NUMBER 2 101 


little larger when mature. Typical developmental times are shown on 
Table 1. Adults emerged between 31 May and 6 June. 

King’s hairstreak is probably limited to Symplocos tinctoria 
throughout its range, as it was also reported on this plant in Georgia 
(Floyd, 1974), and its distribution agrees well with that of the plant. 
The report by Harris (1972) that flame azalea (Rhododendron calen- 
dulaceum) is a host for this butterfly may have resulted from a mis- 
identified S. liparops. 


Calycopis cecrops 


The habits of Calycopis cecrops are bizarre. Found on both Hat- 
teras and Roanoke Islands, the butterfly has two full broods and a 
partial third (April, July, September). Robert Cavanaugh of North Car- 
olina observed female C. cecrops ovipositing on dead leaves on the 
ground under plants (pers. comm.). With this cue the senior author 
began to watch females in the field and eventually saw the act re- 
peated. The eggs are laid on the underside of dead leaves and are 
hidden from view. The one found by Gifford was laid on a dead leaf 
three inches from the base of the nearest plant, a wax myrtle (Myrica 
cerifera). The larvae were dull blackish brown and densely setate in 
all their instars. 

On 8 August 1980, several female Calycopis were placed in a cage 
with sprigs of wax myrtle and staghorn sumac (Rhus typhina), as well 
as some fall flowers. Small dead dry leaves were scattered on the 
bottom of the cage. The females laid a number of eggs, all under the 
dead leaves hidden from view. When the eggs hatched, the young 
larvae had their choice between wax myrtle and sumac. Wax myrtle 
was selected by 36 larvae (59%) and sumac by 25 (41%). 

Fresh leaves of the plant selected were provided daily, and all lar- 
vae survived until hibernation in late fall. Larvae on sumac grew at 
a normal rate, although their growth slowed after the third molt. The 
molting process normally required about three days, but was length- 
ened to five or six days for the third molt. After this molt the larvae 
each rested under a sumac leaf until the following March when they 
pupated. Sixteen larvae (64%) survived until spring. 

The larvae on wax myrtle grew more slowly but molted at the same 
time as the sumac group. They were relatively smaller and spent more 
time resting. The three larvae (8%) which successfully overwintered 
began moving about with the first warm weather in February. The 
larvae were then provided wax myrtle sprigs with overwintered leaves 
and male flower buds. The larvae fed on the flower buds and ignored 


102 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


the leaves. They completed development and pupated about the time 
the flowers opened in nature. 

Wax myrtle is the primary natural host in coastal North Carolina, 
although sumac and oak may be selected on occasion. The fact that 
different hosts may be used results in different developmental times 
and, therefore, extended flight periods. For example, spring genera- 
tion adults may be found from 10 April to 15 May. 

The life stages of this species were previously described by Rawson 
& Hessel (1951). They associated the butterfly with dwarf sumac (Rhus 
copallina) in New Jersey, and obtained eggs by enclosing females in 
paper bags with suspected host material. They reported the larvae to 
prefer leaves and to show strong negative phototropism. 


Mitoura hesseli 


On 20 July 1980, three female M. hesseli were collected near East 
Lake, Dare County, North Carolina, visiting flowers of sweet pep- 
perbush (Clethra alnifolia). In the vicinity grew white cedar (Cham- 
aecyparis thyoides), the only host of C. hesseli. The females were 
taken back to Roanoke Island where they were confined with branch- 
es of white cedar and flowering branches of sweet pepperbush. One 
female died the first day, but the other two lived until about 25 July 
by which time 20 eggs had been laid. 

The larvae were transferred to red cedar (Juniperus. BH a 
since white cedar was not readily available. Two larvae would not 
accept the alternate host and died without feeding. The remaining 
larvae accepted red cedar and developed normally. The young larvae 
were bright green. At the third molt the white transverse markings 
began to appear, but were not fully developed until the last instar. 
Larvae in the third instar would hang by a silk thread when acciden- 
tally dislodged. Some larvae molted five times prior to pupuation and 
others molted only four times. M. hesseli larvae required two days to 
molt before feeding was reinitiated. Larvae always ate the shed exu- 
viae. 

The life history of Mitoura hesseli was described previously by 
Rawson et al. (1951), and the observations reported herein are gen- 
erally confirmatory. This hairstreak is well known to be closely as- 
sociated with its host white cedar throughout its range from New 
Hampshire to Florida. 

Previously, Remington and Pease (1955) found that Mitoura gry- 
neus larvae, whose normal host is red cedar (Juniperus virginiana), 
will readily feed and develop normally on white cedar. Thus, results 
of the reverse experiment reported here are of more than passing 
interest. 


VOLUME 37, NUMBER 2 103 


Incisalia henrici 


On Hatteras Island, I. henrici fed only upon yaupon (Ilex vomito- 
ria), but on Roanoke Island, only American holly (Ilex opaca) was 
utilized. Individuals of the Hatteras Island population had a greenish 
cast ventrally, and might eventually be described as a separate sub- 
species (L. D. Miller, pers. comm.). 

Female I. henrici oviposited when the hollies were beginning to 
leaf out. Females circled about investigating expanding buds. If a bud 
seemed suitable a female would fly to a nearby leaf (the previous 
year s—holly being an evergreen) and laid an egg on the upper sur- 
face adjacent to the midrib. Eggs were laid indiscriminately with re- 
gard to sex of the host (Ilex opaca being dioecious). Eggs hatched 
seven to 11] days after oviposition. Upon hatching the young larvae 
crawled to the expanding leaf buds and bored in. As the leaves ex- 
panded, the larvae skeletonized them; then as they became larger, 
they fed on entire leaves. Fruits or flowers were not fed upon. Older 
larvae fed at night and rested under year-old leaves by day. 

I. henrici larvae were green, molted three times, and required about 
a month for development to pupation. 

On Hatteras Island, adult I. henrici fed at willow flowers. Typical 
developmental times are shown on Table 1. Adults emerged from 10 
to 16 April. 

The geographic distribution of I. henrici’s host associations are un- 
usual. The butterfly, which uses a single host in any one area, feeds 
on quite different hosts in different portions of its range. In much of 
its inland range, the insect selects redbud (Cercis canadensis) with 
its larvae feeding on flowers and young fruits (many reports). Appar- 
ently, Atlantic coastal plain populations are associated with various 
hollies, as there are also reports from Florida (Baggett, 1980) and New 
Jersey (W. Wright, pers. comm.). In Texas, henrici feeds on persim- 
mon (Diospyros texana) (Kendall, 1964), while the larva feeds on 
huckleberry (Gaylussacia) or blueberry (Vaccinium) on the south- 
eastern Piedmont and in the upper Great Lakes States (Harris, 1972; 
Baggett, 1980; Nielsen, 1970). Shapiro (1966) reports it on wild plum 
(Prunus) and possibly blueberry in western Pennsylvania. 


Fixsenia ontario 


In nature, females must usually oviposit on high branches of their 
host plants, since larvae were never found on low branches. In ad- 
dition, females accepted as oviposition substrata only twigs which 
would produce male catkins the following spring. On Hatteras Island, 
a female was observed depositing an egg on laurel oak (Quercus laur- 
ifolia). She was taken alive and caged with branches of both laurel 


104 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


oak and live oak, the only oaks found at that locality. The ontario 
female laid six eggs on the former and eight on the latter. On Hatteras, 
the host leaves began to expand before the eggs hatched, and larvae 
always ate host leaves. There was no difference in rate of growth, size 
or timing of adult emergence for larvae raised on the two hosts. 

On Roanoke Island, matters were quite different; two females caged 
with branches from non-reproductive individuals of QO. laurifolia laid 
only two or three eggs before dying. The following spring the larvae 
hatched before the hosts had begun to leaf out. At that time both oaks 
were in flower, so the larvae were provided with male catkins, which 
they fed upon until young leaves were available. The following year 
females were caged with branches of reproductive-aged trees of the 
hosts mentioned above, as well as black jack oak (Quercus marilan- 
dica), blue jack oak, and Spanish oak (Q. falcata). All five occurred 
naturally on that island. All females were caged together, and quite 
a lot of eggs were laid but only on Q. virginiana. Subsequent attempts 
to obtain oviposition on black jack, blue jack and Spanish oaks were 
also unsuccessful. A few larvae obtained from eggs laid on live oak 
were fed catkins and leaves of black jack oak. They developed nor- 
mally and adults eclosed at the same time as those raised on live oak. 

Young larvae bored into individual flowers and fed on pollen leav- 
ing the outer portion uneaten. The catkins were available until the 
third larval molt by which time the young leaves appeared. The larvae 
finished their feeding on young leaves. The larvae were always pale 
green, about the shade of the lower surface of a live oak leaf. In nature, 
they must pupate on the leaves or branches. In captivity, they usually 
pupated on leaves on the bottom of the container. Developmental 
times for a typical individual are given on Table 1. Adults emerged 
from 22 May to 6 June. 

Little has been published previously on the life history of this in- 
sect, save for a few reports of host associations. Kendall (1964) reared 
the butterfly from larvae found on live and laurel oaks in Texas, while 
Shapiro (1974) reported white oak (Quercus alba) as the host in New 
Jersey. Clench (1971) found this hairstreak on shale barrens in Vir- 
ginia and Pennsylvania where bear oak (Quercus ilicifolia) is preva- 
lent. 


ACKNOWLEDGMENTS 


We thank Robert K. Robbins and an anonymous reviewer, both of whom read the 
manuscript and provided suggestions for its improvement. 


VOLUME 37, NUMBER 2 105 


LITERATURE CITED 


BAGGETT, H. D. (compiler). 1980. Zone 6. South. 1979 Season Summary. News Lep- 
id. Soc. 1980:21-23. 

BROWN, L. N. 1976. A population of the striped hairstreak, Satyrium liparops lipa- 
rops (Lycaenidae) in west-central Florida. J. Lepid. Soc. 30:213. 

CLENCH, H. K. 1971. Some records of Euristrymon ontario (Lycaenidae). J. Lepid. 
Soc. 25:80-82. 

FLOyD, J.C. 1974. A new foodplant record for Satyrium kingi (Lycaenidae). J. Lepid. 
Soc. 28:353. 

Harris, L., JR. 1972. Butterflies of Georgia. Univ. Oklahoma Press, Norman. 326 pp. 

KENDALL, R. O. 1964. Larval foodplants for twenty-six species of Rhopalocera (Papil- 
ionoidea) from Texas. J. Lepid. Soc. 18:129-157. 

Kxiots, A. B. 1951. A Field Guide to the Butterflies of North America, East of the 
Great Plains. Houghton Mifflin Co., Boston. 349 pp. 

KNUDSEN, J. P. 1955. A new host plant record from Strymon liparops. Lepid. News 
Sala 12. 

NIELSEN, M. C. 1970. New Michigan butterfly records. J. Lepid. Soc. 24:42-47. 

Rawson, G. W. & S.A. HESSEL. 1951. The life history of Strymon cecrops Fabricius 
(Lepidoptera, Lycaenidae). Bull. Brooklyn Entomol. Soc. 46:79-84. 

RAWSON, G.W., J. B. ZIEGLER & S. A. HESSEL. 1952. The immature stages of Mitoura 
hesseli Rawson and Ziegler. Bull. Brooklyn Entomol. Soc. 46:123-130. 

REMINGTON, C. L. & R. W. PEASE. 1955. Studies in foodplant specificity. 1. The 
suitability of swamp white cedar for Mitoura gryneus (Lycaenidae). Lepid. News 
9:46. 

SHAPIRO, A. M. 1966. Butterflies of the Delaware Valley. American Entomol. Soc., 
Spec. Publ. 79 pp. 

1974. Butterflies and skippers of New York State. Search 4:1-60. 

SMITH, J. W. 1797. A Natural History of the Rarer Lepidopterous Insects of Georgia. 
London. 2 vols., 214 pp. 


Journal of the Lepidopterists’ Society 
37(2), 1983, 106-114 


NOTES ON THE SATYRID BUTTERFLY POPULATIONS 
OF CORCOVADO NATIONAL PARK, COSTA RICA 


PAUL L. WHITTAKER 
Department of Zoology, University of Texas, Austin, Texas 78712 


ABSTRACT. Results of a mark-release recapture study of satyrid butterflies at Cor- 
covado National Park, Costa Rica are reported. Seven species were captured in undis- 
turbed primary forest along the Olla Trail. Eleven species, including all but one of the 
species from the undisturbed habitat, were found in the more heterogeneous habitat 
along the Rio Claro Trail. Two of the principal forest species, Pierella luna Fabricius 
and Euptychia arnaea Fabricius, occur in “viscous” populations (repeated recaptures 
within a relatively small area), with a predominance of males among captured individ- 
uals. The population structure of Cithaerias (Callitaera) menander Drury was ex- 
tremely “fluid” (no recaptures), with mostly females captured and no behavioral pref- 
erence for local high points. The other major understory species, Pierella helvetia 
incanescens Godman & Salvin, was not caputred often enough to make any significant 
biological inferences. Results are discussed with reference to theories of competition, 
character displacement and ecological disturbance. 


The butterfly families Satyridae, Morphidae and Brassolidae have 
been called a “family cluster’ by Young & Muyshondt (1975). They 
are characterized by monocotyledonous larval food plants (although 
many morphos feed on legumes), use of non-floral resources (fallen 
fruit, dung, etc.) as adults (many north-temperate satyrids do feed on 
flowers) and cryptic, brown to ochre coloration (irridescent blue on 
the upper wing surface is fairly common among tropical species). 
Monocots have relatively few secondary compounds, and the gener- 
ally cryptic color patterns of adults may result from inability to se- 
quester toxic compounds. Larval food preference among monocot 
feeders does not appear to have “coevolved” along with plant de- 
fenses as it has in other groups (Ehrlich & Raven, 1964; Gil- 
bert, 1975). Tropical Euptychia species (used here and afterwards 
in the sense of Weymer, 1924) commonly accept several species 
of grasses as larval hosts (M. C. Singer, pers. comm.). Young & Muy- 
shondt (1975) report two congeneric species of brassolids which use 
larval hosts in different families, and in another paper (Young & Muy- 
shondt, 1972) emphasize the importance of factors such as habitat 
selection in explaining radiations within the genus Morpho. 

Satyrids, like many groups of new world organisms, reach their 
greatest diversity in the tropics. Ecology of tropical satyrids has re- 
ceived little attention to date, with the exception of work by Young 
(1972) and Singer (unpublished). Young (1972) marked individuals of 
twelve species (including satyrids, brassolids and morphids) at four 
Coumarouna oleifera (Leguminosae) fruit drops at La Selva, Costa 
Rica. He found little day to day variation in numbers, with the same 


VOLUME 37, NUMBER 2 107 


butterflies returning repeatedly to the same feeding sites. Adult mor- 
tality and recruitment were low, and the species composition at the 
four fruit drops was essentially homogeneous. Most of the species 
used artificial food when it was offered, but introduction of artificial 
food did not result in separation of feeding preferences, as would be 
expected if competition for adult resources were a limiting factor. This 
study presents basic data on satyrid distributions over a variety of 
habitats and behavioral comparisons of several species which occupy 
the same habitat and use the same adult food resources. 


METHODS 


Corcovado National Park is located on the Osa Peninsula of Costa 
Rica on the southern Pacific coast. The park headquarters is located 
in a valley about 1 km from the ocean. The lowlands near headquar- 
ters have been used for agriculture in the past, and several habitat 
types are found, including pasture, second growth and disturbed for- 
est. The surrounding hills are covered with tropical wet forest. This 
study was carried out in July, which is early in the wet season. Two 
censusing routes were used, one (the Rio Claro Trail) to sample dis- 
turbed habitats and another (the Olla Trail) to sample primary forest. 
A map of the Sirena (headquarters) area of Corcovado National Park 
is included in Gilbert et al. (in prep.). The Rio Claro Trail starts just 
north of headquarters, goes southward up a steep ridge through dis- 
turbed primary forest, bears to the north and forks about 0.8 km (one- 
half mile) from its beginning. The left fork goes to a small landslide 
clearing and then ends. The right fork goes south along a long ridge, 
with the Rio Claro to the north. Vegetation along the ridge is very 
heterogeneous, with Heliconia (Musaceae) and Cecropia (Moraceae) 
intermixed with the primary forest species. The trail descends the 
ridge through an extensive stand of Heliconia, goes past a ruined 
shack in a small clearing, then turns and goes west along the Rio Claro 
until it reaches the ocean. The forest along the river contains a higher 
proportion of mesophytic species because of the level topography and 
consequently increased soil moisture. Total walking distance is about 
3.5 km. The Olla Trail goes northeast through slightly disturbed to 
undisturbed upland forest, after crossing the Rio Camaronal near 
headquarters. The first 2.5 km were used for study. Both trails were 
marked with metal tags on tree trunks at intervals of about 50 m. 

Censusing was done by walking along the trail and catching as 
many satyrids as possible. Routes were usually run in the morning 
between 0730 h and noon. Afternoon rain was very common, and 
butterfly activity along the trails was somewhat less during the after- 


108 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


NUMBER OF INDIVIDUALS CAPTURED 
< N 


Ol 
N 


= 
7] // / 7 CIIHAERIAS (CALLITERA) | MENANDER DRURY 


777 | (PIERELLA ) (PIERELLA HELVETIA INCANESCENS GODM. & SALV) 
\) TAYGETIS ANDROMEDA CRAMER 

() EUPTYCHIA LIBYE LINNEAUS 

i EUPTYCHIA INSOLATA 


Fic. 1. Total number of butterfly individuals captured along the Olla Trail. 


noon. The Rio Claro Trail was sampled on 6, 7, 8, 9, 10, 14, 19, 21, 
27 (afternoon) and 28 July 1979, usually starting at the mouth of the 
river. The Olla Trail was sampled on 11, 12, 13, 15, 18, 20, 25, 27 and 
29 July. Capture success was about 60% for most species and some- 
what lower for Pierella helvetia incanescens Godman & Salvin. 

Captured butterflies were marked with a red PILOT ultrafine point 
pen and then released. (A few butterflies were retained for voucher 
specimens.) All butterflies were numbered sequentially, regardless of 
species. The number was written on the undersurface of one hind- 
wing and opposite forewing. Number, sex, species, condition (fresh, 
worn or intermediate), date, time and location (measured by pacing 
to the nearest metal marker) were recorded for each capture. Recap- 
tures were released promptly after noting the number and condition. 

Horizontal distance between markers was estimated by pacing along 
the trail, using a compass to determine direction. Capture points were 
marked on a map, and distances between consecutive capture points 
for recaptured butterflies were estimated from the map distances. 
These estimates do not include vertical displacement. Each marker 
station was rated for local topography (“0” for level ground or local 
depression, “+” for a slope, “++” for a ridge or hilltop). Abundance 
of fallen fruit in the vicinity of the marker (“0’, none visible; “1”, 
scarce; “2”, moderately abundant; “3’’, abundant) was recorded on 20 
July. There was some fluctuation in the abundance of fallen fruit over 
the course of the study, and no attempt was made to identify fallen 
fruit to species. Chi-square tests were used to evaluate the signifi- 
cance of observed trends. 


VOLUME 37, NUMBER 2 109 


NUMBER OF INDIVIDUALS CAPTURED 
a NO 


CN) 
4N NO 


ANWEZc=TSEANNNANNNANN 
\\\\\\\\ Jj EUPLYCHIA HERMES SOSYBIUS 
Kel EUPTYCHIA LIBYE 


TAYGE TIS ANDROME DA 
CITHAERIAS (CALLITAERA) MENANDER 
EUPTYCHIA CONFUSA 


Fic. 2. Total captures along the Rio Claro Trail. 


RESULTS 


Figs. 1 and 2 show the number of different butterflies of each species 
captured in each of the two study areas, ranked in order of the number 
of individuals captured. The Rio Claro Trail fauna is more diverse (11 
species, 1/(% p,?) = 4.40) than that of the Olla Trail (7 species, 
1/(% p;”) = 3.03), in spite of the smaller sample size. Members of the 
related genera Euptychia (Weymer) and Taygetis comprise 59% (39 
out of 66) of the Rio Claro sample and only 22% (18 out of 82) of the 
Olla Trail sample. Several species usually found in open or second 
growth habitat (Taygetis andromeda Cramer, Euptychia hermes Lin- 
naeus, Euptychia usitata Butler, Euptychia hesione Sulzer, Eu- 
ptychia libye Linnaeus) were captured in or near the clearing on the 
Rio Claro Trail. Cithaerias menander Drury was much less abun- 
dant in the Rio Claro area (n = 2 vs. n = 20). 

Table 1 shows basic capture data for the four most common Olla 
Trail satyrids. Pierella luna Fabricius and Euptychia arnaea Fabri- 
cius were predominantly males. Recaptures were common in both 
species. One P. luna was captured six times within a 5 m radius, while 
another moved at least 1268 m between four capture points. The long- 
est movement between captures by any E. arnaea was 30 m. Most of 


110 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 1. Capture-recapture data on major Olla Trail satyrid species. 


Mean recapture 
No. of but- No. No. re- No. of re- distance (S.D.) 
terflies sampled captured captures (m) 


Pierella luna 3 35 0 DD 48 73.2 (107.2) 
2 5 2 0 0 ie 
Cithaerias menander } 6 0) 0 0 eae 
2 14 9) 0 0 as 
Euptychia arnaea fe) 14 0 5 8 7.6 (12.6) 
g 1 0 0 0 ae 
Pierella h. incanescens fe) il 0 0) 0 = 
2 3 i 0 0 = 


the C. menander captures were females, and there were no recaptures 
of this species. The biased sex ratios are probably a result of behav- 
ioral differences between the sexes. The absence of C. menander 
recaptures indicates a large effective population size, although the 
density per unit area for C. menander and P. luna may be roughly 
comparable. 

Table 2 gives the number of male P. luna captured (Olla Trail), the 
number recaptured (second and subsequent captures of the same but- 
terfly on the same day are omitted), and a simple Lincoln index (N = 
n(C + R)/R, where n is the number of individuals previously marked, 
N is the estimated population density, C is the number of new cap- 
tures on that day and R is the number of recaptures) estimate of pop- 
ulation number for each day of the study. The mean estimated pop- 
ulation size is 41.85, with standard deviation 13.96. By sampling along 
the trail, I believe I effectively covered an area of about one hectare. 

Table 3 shows the number of captures in relation to topography. 
There were six level stations (“0”) covering 367 m of trail, 21 stations 
on slopes (“+’’, 1201 m) and 17 stations on ridges or hilltops (“++”, 
914 m). Expected values were calculated by multiplying the number 
of total captures (including recaptures) for each species by the pro- 


TABLE 2. Number of new captures (C), recaptures (R) and Lincoln-index estimates 
of population density (N) of P. luna males on the Olla Trail. 


Date (July) 
ll 12 13 15 18 20 25 ‘27 29 
C 9 6 3 4 U 3 1 1 1 
R 0) DY) 4 ©) 3 3) th 11 8 
N 36.0 26-29 ADO Tedd) 46.4 36.6 36.0 38.25 


VOLUME 37, NUMBER 2 lll 


TABLE 3. Total captures (Olla Trail), by topographic position of nearest marker. 
Expected values in parentheses. See text for further explanation. 


0 + ++ 

P. luna 2 4] 45 Yo — 426 dis— 2 P<10:005 
(Qin (42158)—" (32741) 

C. menander 0 Te 9 9 = S00), Gli = 2 IP > O10 
(2.96) (9.68) (7.37) 

E. arnaea 0 7 16 == ID. che = BP 0S 
(3.40) CE 13) (8.47) 

P. h. incanescens 0 0 4 P = 0.0184 


portion of trail belonging to that given topographic category. For ex- 
ample, the expected number of P. luna captures at level stations is 
(88)[367/(367 + 1201 + 914)]. P. luna and E. arnaea both had a sig- 
nificant preference for local high points. C. menander captures showed 
no significant trend. P. h. incanescens also seemed to prefer high 
points, although the number of captures was very small. 

Table 4 shows total captures for each species, according to local 
abundance of fallen fruit on 20 July. Two stations (122 m of trail) had 
no fruit visible nearby, 26 stations (1378 m) were rated as “1”, 11 
stations (655 m) were rated as “2” (moderately abundant) and five 
stations (327 m) were rated as “3”. Expected values were calculated 
by multiplying the total number of captures by the proportion of trail 
assigned to that rating group. Capture distributions were significantly 
non-random for all species except P. h. incanescens. For the other 
three species, there was a trend toward higher capture frequencies 
around high densities of fallen fruit. The trend is less pronounced for 
P. luna than for C. menander or E. arnaea. Eighteen of the 23 E. 
arnaea captures were within 100 m of a single large fruit drop. The 


TABLE 4. Total captures (expected values in parentheses) by abundance of fallen 
fruit around the nearest marker (Olla Trail). 


Fruit abundance 


0 1 2 3 
P. luna 2 39 34 13 Me 16-402 dt — oa be—a0205 
(4.31) (48.89) (23.21) (11.59) 
C. menander 0 i 3 10 x” = 20.08, df = 3, P < 0.005 
(O98) (TEI PS 28)) 42268) 
E. arnaea 0) 1 il 10 Yo Adi — 3 re 0005 


GIES) Gl 257.8) wee (GOW) me (S-03) 
P. h. incanescens ) 2 1 ] 


112 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 5. Total captures (Olla Trail) by time of day. 


Time 
0730— 0830— 0930- 1030— 1130- After 


0830 0930 1030 1130 1230 8 1230 


P. luna 14 24 26 18 4 2; 
C. menander 8 5 8 2 ] 1 
E. arnaea Il 118} 5 3 1 0 
P. h. incanescens 0) ] ] 1 ik 0 


C. menander Ptures were distributed among several widely separated 
sites. There was no significant association between topography and 
abundance of fallen fruit. 

Table 5 shows the number of captures by time of day for P. luna, 
C. menander, E. arnaea and P. h. incanescens. C. menander were 
most active early in the morning. E. arnaea were most active around 
0900 h, and P. luna showed a slight peak around 1000 h. Very little 
time was spent collecting after 1200 h, so the picture presented in 
Table 5 is incomplete. Wet weather usually depressed activity in late 
morning or early afternoon, but all four species were seen flying on 
sunny afternoons. 


DISCUSSION AND SUMMARY 


Cutting along the Rio Claro Trail has created openings and allowed 
invasions by successional plant species, such as Heliconia. The great- 
er habitat diversity results in greater satyrid diversity, with several 
Euptychia species occurring which are not found in the forest. This 
result supports recent suggestions of Gilbert (1980) and Huston (1979) 
that ecological disturbance may increase species diversity. 

The behavior of C. menander reported here is markedly different 
from that described by Young (1972). Young reported small, closed 
demes, with the same individuals returning to the same place day 
after day. The opposite seems to be true at Corcovado: Different in- 
dividuals were captured at the same place and time on different days. 
Young (1972) also found a mid-day peak in activity, with no sightings 
before 0830 h. Several hypotheses may be proposed as explanations. 
The habitat at Corcovado is more hilly, so fruit drops might be smaller 
and less persistent through time. This would generate a more fluid 
population structure. Repeated movements in the same area might 
increase chances of predation, although there is no reason to believe 
this should be more important at Corcovado. A more scattered, 
ephemeral distribution of larval resources at Corcovado might also 
result in a more mobile population. Interference competition with P. 


VOLUME 37, NUMBER 2 113 


luna may also play a role. P. luna (which is not present at Young’s 
study site) is an aggressively territorial species and often chases con- 
specifics. The early morning activity peak and mobile flight behavior 
of C. menander would both reduce the chances of being chased by a 
P. luna. Young (1973) reports aggressive patrolling behavior in one of 
his La Selva species (Morpho amathonte Deyrolle). According to 
Young (1972), M. amathonte flies early in the morning at La Selva 
before C. menander becomes active there. 

P. luna usually flies low to the ground, often following the contour 
of the trail for 20 or 30 m. They are extremely cryptic against the 
background of forest litter: Young individuals resemble wet leaves 
when at rest, and older butterflies resemble dry leaves. Males will 
patrol the same area repeatedly, and encounters often result in chas- 
ing behavior. Positioning on ridges and hilltops may confer some ad- 
vantage in mating, as Shields (1967) has reported in several temperate 
species. According to Scott (1968), “hilltopping” is usually an adap- 
tation to low population densities, but densities of P. luna were not 
especially low during the study. Access to food may also be important. 
Occasional long flights may reflect searching for a territory with ad- 
equate food and few competitors. 

E. arnaea is usually encountered in small sunny areas within the 
forest, with several males present in the same place. Scott (1974) lists 
several situations: which should favor “perching” (as opposed to pa- 
trolling) as a mate location strategy, including low population density 
and patchy resource distribution. E. arnaea is less common and more 
patchily distributed than P. luna at Corcovado, so either one could 
be responsible for the observed behavior. Both predominantly male 
species (P. luna and E. arnaea) prefer local high points, while C. 
menander, which flies singly (usually female) or in pairs at Corcovado, 
doesn't. Cithaerias and Pierella are placed close together in most 
classification schemes (Weymer, 1924), even though their behavior in 
this situation is very dissimilar. Both C. menander and E. arnaea 
show slight sexual dimorphism with respect to size (males are smaller) 
and wing color. P. luna males have conspicuous scent patches (an- 
droconia) near the upper rear edge of the hindwings. 

Previous work by Gilber & Singer (1973) has demonstrated that 
spatial distribution of resources may affect flight and mating behavior 
in butterflies. Results presented here indicate that interspecific com- 
petition may also have a significant impact. Comparison of behavior 
of related species in the same area and of single species in different 
areas, combined with accurate measurement of niche parameters such 
as food preference and flight time, will be needed to clarify this issue. 
Detailed descriptions of courtship, mating, oviposition behavior and 
larval life histories are also necessary. 


114 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


ACKNOWLEDGMENTS 


I thank L. E. Gilbert and J. Mallet for their interest and encouragement, and the staff 
of Corcovado National Park for their hospitality during my stay. P. J. DeVries and M. 
C. Singer helped identify specimens. This research was supported in part by a graduate 
student research grant from the University of Texas at Austin. 


LITERATURE CITED 


EHRLICH, P. R. & P. F. RAVEN. 1964. Butterflies and plants: a study in coevolution. 
Evolution 18:586-608. 

GILBERT, L. E. 1975. Ecological consequences of coevolved mutualism between 
butterflies and plants. Pp. 210-240, in L. E. Gilbert & P. H. Raven (eds.). Coevo- 
lution of Animals and Plants. University of Texas Press, Austin. 246 pp. 

1980. Food web organization and the conservation of tropical diversity. Pp. 
11-33, in M. Soule & B. Wilcox (eds.). Conservation Biology. Sinauer Assoc., Sun- 
derland, Mass. 

GILBERT, L. E. & M. C. SINGER. 1973. Dispersal and gene flow in a butterfly species. 
American Nat. 107:58—72. 

Huston, M. 1979. Toward a general hypothesis of species diversity. American Nat. 
113:81-101. 

Scott, J. A. 1968. Hilltopping as a mating mechanism to aid the survival of low 
density species. J. Res. Lepid. 7:191—204. 

1974. Mate locating behavior in butterflies. American Mid. Nat. 91:103-117. 

SHIELDS, O. 1967. Hilltopping. J. Res. Lepid. 6:69-78. 

WEYMER, G. 1924. Satyridae. Pp. 193-183, in A. Seitz (ed.). Macrolepidoptera of the 
World. Vol. V. The American Rhopalocera. Alfred Verlag, Stuttgart. 1139 pp. 
YOUNG, A.M. 1972. Community ecology of some tropical rain forest butterflies. Amer- 

ican Mid. Nat. 87:146—157. 

1973. Studies on comparative ecology and ethology of several species of 
Morpho butterflies. Studies Neotrop. Fauna 8:17-50. 

YOUNG, A. M. & A. MuYSHONDT. 1972. Geographical and ecological expansion in 
tropical butterflies of the genus Morpho in evolutionary time. Rev. Biol. Trop., 
Costa Rica 20:231-263. 

1975. Studies on the natural history of Central American butterflies in the 

family cluster Satyridae-Brassolidae-Morphidae. III. Opsiphanes tamarindi and 

Opsiphanes cassina in Costa Rica and E] Salvador. Studies Neotrop. Fauna 10:19- 

56. 


Journal of the Lepidopterists’ Society 
37(2), 1983, 115-120 


A NEW SUBSPECIES OF SPEYERIA EGLEIS 
(NYMPHALIDAE) FROM THE PUMICE REGION 
OF CENTRAL OREGON 


PAUL C. HAMMOND AND ERNST J. DORNFELD 
Department of Zoology, Oregon State University, Corvallis, Oregon 97331 


ABSTRACT. A small-sized Speyeria egleis of central Oregon, delimited in its dis- 
tribution to an area of some 4026 km? (2500 square mi.) extending northeasterly of 
Crater Lake, is described as a new subspecies, S. e. moecki. This region is geologically 
characterized as one of heavy ash and pumice outfall from the eruption of Mt. Mazama. 
The new subspecies is distinguished from the more southerly and larger S. e. oweni 
and from the sympatric and superficially similar S$. mormonia erinna. 


Lepidopterists of the Pacific Northwest have long been aware of a 
distinctive population of Speyeria egleis (Behr) in the pumice region 
of central Oregon (Deschutes and Klamath Counties). This was first 
noted by Arthur H. Moeck (1957), who took specimens in the Sand 
Creek region just east of Crater Lake at about 1524 m (5000 ft) ele- 
vation. Unofficially known as the Sand Creek type, this small-sized 
egleis was recently illustrated by Dornfeld (1980, Pl. 30, Figs. 3 and 
4). Remarkably uniform in phenotype, it is geographically confined 
to an area of some 4026 km? (2500 square mi.), as shown in the dis- 
tribution map (Fig. 1). This region is geologically characterized by 
heavy ash and pumice outfall from the eruption of Mt. Mazama (Crater 
Lake) about 6600 years ago. Tilden (1963) has given an account of the 
geological history and ecology of the Sand Creek Basin, but with 
respect to Argynnis (=Speyeria) he believed that “no equilibrium in 
phenotype has been reached by any of the several species.” The evi- 
dence as it affects S. egleis does not sustain this view. Both with 
respect to the relative uniformity of its distinctive phenotype and the 
circumscription of its distribution, the “Sand Creek” egleis can be 
readily separated from the egleis populations that occur in southern 
Jackson and Klamath Counties. Those populations are composed of 
consistently larger butterflies that show affinities with S. e. oweni 
(Edwards) of the Mt. Shasta region. The Sand Creek egleis is here 
recognized as a distinct subspecies. Several hundred specimens have 
been examined. 


Speyeria egleis moecki, new subspecies 


Male. Length of forewing (n = 71) 21 to 25 mm (x = 23 mm). Dorsal wing surfaces 
medium orange with the usual pattern of black spots and bars. Moderate basal suffusion 
and veins of forewing slightly thickened with dark scales. Dorsal hindwing with yel- 
lowish patches between the black median band and black postmedian spots. Ventral 
forewing with yellowish or yellow-orange ground color; brown patches around the 


116 JOURNAL OF THE’LEPIDOPTERISTS’ SOCIETY 


Deschutes Co. 


Douglas Co. 


Lake Co. 


Klamath Co. 


Jackson Co. 


Fic. 1. Distribution of Speyeria egleis moecki, n. ssp. Figures in circles identify 
collection sites. 


postmedian spots of the apical region. Ventral hindwing with dark reddish brown or 
umber-brown disc and a narrow yellowish tan or pale brown submarginal band. Spots 
usually brightly silvered and narrowly outlined basally with black scales. Median spots 
usually small, rounded or pointed, and the submarginal spots rounded to flattened. 
Brown or reddish brown marginal border. 


VOLUME 37, NUMBER 2 SLL 


Fic. 2. Speyeria egleis moecki, n. ssp. Above: dorsal and ventral sides of holotype, 
male. Below: dorsal and ventral sides of allotype, female. 


Female. Length of forewing (n = 54) 22 to 27 mm (x = 25 mm). Similar to the male, 
but veins of forewing not thickened; dark basal suffusion usually more extensive. Ven- 
tral forewing strongly flushed with reddish orange toward base of wings, but pale 
yellow along costal margin (as in male). 

The name chosen for this race of Speyeria egleis honors the memory of the late 
Arthur H. Moeck of Milwaukee, Wisconsin. Over a period of thirty summers he and 
his wife Dorothy systematically covered the North American Continent in search of 
Fritillaries and, thereby, contributed greatly to our knowledge of these butterflies. 

Types. HOLOTYPE: male (Fig. 2), Skookum Mdw., Walker Rim, Klamath Co., Ore., 
25 July 81 (P. C. Hammond). ALLOTYPE: female (Fig. 2), same data. Types deposited 
in the American Museum of Natural History. PARATYPES: 48 males and 33 females. 
Disposition as follows: one pair each to the U.S. National Museum of Natural History, 
the California Academy of Sciences, and the Systematic Entomology Laboratory of 
Oregon State University; 8 males and 4 females to the collection of L. Paul Grey; 17 
males and 10 females retained by Paul C. Hammond; 20 males and 16 females retained 
by Ernst J. Dornfeld. 

Records. Figures in parentheses identify numbered loci on map (Fig. 1). DES- 
CHUTES CoO.: (1) Three Creeks Mdw., 6/viii/81 (Hammond); (2) Davis Lake, 24/vii/ 
34 (Jewett); (3) 6 mi. E La Pine, 16/vii/61 (Shields); (4) Paulina/East Lakes, 15/vii/59 
(Moeck), 16/vii/61 (Shields), 27/vii/75 (Hammond). KLAMATH CoO.: (5) Crescent Lake, 
2.3/vii/60 (Newcomer); (6, 7) Gilchrist, Crescent, 29/vii/45, 10/vii/54, 4/vii/56, 28/vii/57, 
4/vii/59, 10/vii/60, 2/vii/67 (Domfeld); (8) Mowich, 11/vii/54 (Domfeld); (9) Round Mdw., 
22/vii/68 (Hinchliff); (10) Cannon Well, 30/vii/61 (Newcomer); (11) South Walker Spg., 
22/vii/68 (Hinchliff); (12) Skookum Spg., 23/vii/61 (Newcomer); (13) Skookum Mdw., 
23/vii/61 (Newcomer), 17/viii/62, 13/viii/64, 18/vii/66 (Dornfeld), 24/vii/66, 18/vii/69 
(Hinchliff), 17/vii/79 (Lattin), 26/vii/75, 25/vii/81 (Hammond); (14) Dempsey Spg., 18/ 


118 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Fic. 3. Top: Speyeria egleis oweni, male, dorsal and ventral sides; Mt. Shasta, 
Siskiyou Co., Calif., 23 July 81 (PCH). Middle: S. egleis moecki, male, dorsal and 
ventral sides; Sand Cr. at Hwy. 232, Klamath Co., Ore., 3 July 68 (EJD). Bottom: S. 
mormonia erinna, male, dorsal and ventral sides; Skookum Mdw., Walker Rim, Klamath 
Co., Ore., 24 Aug 72 (EJD). 


vii/66 (Dornfeld), 25/vii/81 (Hammond); (15) Huckleberry Spg., 24/vii/66 (Hinchliff), 
25/vii/81 (Hammond); (16) Davis Flat, 17/viii/62 (Newcomer), 25/vii/81 (Hammond); 
(17) Beaver Marsh, 17/viii/62 (Newcomer); (18) 5 mi. E Beaver Marsh, 14/vii/61 (Shields), 
13/viii/64 (Dormfeld), 24/vii/66 (Hinchliff); (19) Crater Lake, 8/viii/30, 14/viii/30 (Scul- 
len); (20) Sand Cr. at Hwy. 232, 16/vii/55 (Moeck), 10/vii/62, 25/vii/62, 12/viii/64, 3/vii/ 
68, 20/vi/78 (Dornfeld); (21) Sand Cr. nr. Chinchalo, 22/vii/68 (Hinchliff); (22) 3 mi. E 
Klamath Fst. Nat. WIf. Refuge, 25/vii/81 (Hammond); (23) N of Little Yamsay Mt., 27/ 
vii/64 (Perkins). 


VOLUME 37, NUMBER 2 119 


DISCUSSION 


The only race of Speyeria egleis that comes geographically close to 
the newly described subspecies is S. e. oweni (Edwards), whose type 
locality is Mt. Shasta in Siskiyou County, California (Fig. 3). Popu- 
lations of the oweni phenotype extend northward into the Cascades 
of southern Jackson and Klamath Counties, Oregon, but lie south of 
the region occupied by S. e. moecki. Those populations that extend 
between Lake-of-the-Woods in southern Klamath County and the south 
edge of Crater Lake National Park exhibit clinal intergradation be- 
tween oweni and moecki with respect to size and coloration. How- 
ever, the oweni phenotype is almost completely excluded from the 
moecki populations of northern Klamath and Deschutes Counties, the 
region of the heavy Mazama ash fall (Fig. 1). 

Speyeria egleis moecki can be readily distinguished from the race 
oweni by its uniformly small size, reduced dark basal suffusion, rel- 
atively thinner veins of the male dorsal forewing, and the high fre- 
quency of a reddish brown disc color on the ventral hindwing. For 
comparison, a sample of Mt. Shasta oweni included 85 males with a 
forewing length of 24 to 28 mm (x = 26 mm) and 34 females with a 
forewing length of 26 to 30 mm (x = 28 mm). The sympatric S. mor- 
monia erinna (Edwards) is superficially similar to S. e. moecki in size 
and coloration (Fig. 3), but the latter tends to be darker orange above, 
shows a moderate amount of basal suffusion, and the veins of the male 
are distinctly thickened with dark scales. Speyeria m. erinna, in con- 
trast, is usually pale yellow-orange above, shows almost no basal suf- 
fusion, and the veins of the male forewing are completely thin as in 
the female. In addition, S. m. erinna almost always exhibits a greenish 
tinge along the anal margin of the ventral hindwing that is never 
present in S. e. moecki. Although the adult butterflies of both species 
fly together, the peak flight period of S. egleis usually precedes that 
of S. mormonia by a week or two. 

Speyeria egleis is absent from the Oregon Cascade Range north of 
Deschutes County, but populations of this species do occur eastward 
in the Ochoco Mountains of Crook County. The latter, however, be- 
long to the Rocky Mountain race macdunnoughi (Gunder), which is 
highly divergent from moecki in both size and coloration. The dorsal 
wing surfaces show a very extensive dark basal suffusion, the ventral 
disc color is dark black-brown to greenish brown, and the forewing 
length is 27 to 31 mm in males, 29 to 33 mm in females. Virtually no 
trace of such macdunnoughi influence has been observed in Cascad- 
ian moecki populations, which suggests that moecki is largely derived 
from the adjacent oweni populations of northern California. 


120 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


The distribution of S. e. moecki corresponds very closely with the 
ash-pumice fields deposited by the eruptions of Mt. Mazama, Mt. 
Newberry, and the volcanoes of the Three Sisters system. This ash- 
pumice habitat represents an extremely xeric environment since the 
volcanic material fails to retain moisture during the summer growing 
season. As a result the vegetation is dominated by a shrubby forest of 
lodgepole pine (Pinus contorta), with scattered bitterbrush (Purshia 
tridentata) and small tufts of grasses on the forest floor. However, in 
some areas ground water does come to the surface and produces small 
creeks, seepages, and wet boggy meadows. A great diversity of her- 
baceous plants and butterflies are found in these wet areas. Speyeria 
egleis and its larval foodplant, Viola purpurea (oviposition observed), 
are largely confined to the pine forests adjacent to these habitats. 
Around the upper rim of Crater Lake, however, S. egleis and V. pur- 
purea occupy open, dry, rocky pumice slopes. Speyeria mormonia 
erinna, by contrast, flies in the wet boggy meadows that support Viola 
palustris and V. adunca var. bellidifolia, the larval foodplants of this 
species (oviposition observed). 


ACKNOWLEDGMENTS 


For specimens and data we are obligated to the lepidopterists cited in the distribution 
records. Besides the late Arthur H. Moeck, E. J. Newcomer, and Herman Scullen, these 
include John Hinchliff, Stan Jewett, John D. Lattin, Edwin and Stephen Perkins, and 
Oakley Shields. 


LITERATURE CITED 


DORNFELD, E. J. 1980. The Butterflies of Oregon. Timber Press, Forest Grove, Ore. 
276 pp. 

MoEck, A. H. 1957. Geographic variability in Speyeria. Milwaukee Entom. Soc., 
Special Paper. 48 pp. 

TILDEN, J. W. 1963. The Argynnis populations of the Sand Creek Area, Klamath Co., 
Oregon. Part I. The effect of the formation of Mt. Mazama on the area and its 
possible influence on the butterfly faunas of the Sand Creek Basin. J. Res. Lepid. 
1:109-113. 


Journal of the Lepidopterists’ Society 
37(2), 1983, 121-128 


CAUSAL ANALYSIS OF A MIGRATION OF THE 
SNOUT BUTTERFLY, LIBYTHEANA BACHMANII LARVATA 
(STRECKER) (LIBYTHEIDAE) 7 


RAYMOND W. NECK 


Texas Parks and Wildlife Department, 4200 Smith School Road, 
Austin, Texas 78744 


ABSTRACT. Observation of a massive migration of the snout butterfly, Libytheana 
bachmanii larvata (Strecker), in central Texas in 1971 is described. An association of 
peak migration periods and periodic precipitation episodes is believed to be causal in 
nature. 


One of the many migratory species of Lepidoptera is the snout but- 
terfly, Libytheana bachmanii larvata (Strecker). At irregular intervals 
great numbers of this species migrate in various directions in south 
and central Texas. The latest of these large migrations occurred dur- 
ing July-September 1971. One previous report exists for this migra- 
tion (Helfert, 1972) in which only a portion of the migration is dis- 
cussed. Helfert initiates his remarks with observations on 22 August 
1971 and concludes them on 2 September. Migratory activity was at 
a peak during this period but occurred both before and after this time. 

The purpose of this communication is to supplement the data avail- 
able on this particular migration and to discuss possible environmen- 
tal triggers involved in this phenomenon. A hypothetical relationship 
between snout butterfly population recruitment and periods of sub- 
stantial rainfall is presented. Personal observations of the 1971 flight 
will be presented chronologically before analysis. 

In order to critically analyze a mass migration of snout butterflies 
as the one observed in 1971, one must be aware of normal conditions. 
In central Texas, i.e., the Austin area, snout butterflies are present in 
low numbers, with adults being active on warm winter days (H. G. 
Lacey in Kendall & Kendall, 1971). Despite its constant presence, 
snouts are not apparent to non-lepidopterists, particularly those living 
in urban areas. In south Texas (area south of San Antonio) snout but- 
terflies are often abundant (though highly scattered spatially and tem- 
porally) and quite regularly migrate in large numbers. 

Butterflies of the genus Libytheana are variable in phenotype; 
species relationships are still unclear. The populations of central and 
southern Texas are referred to Libytheana bachmanii larvata (Streck- 
er). Howe (1975) indicated that the taxon of the snout butterflies in 
the Kansas flight (see discussion below) was Libytheana carinenta 
mexicana Michener. This taxon rarely occurs in the United States. 
Clench (1968) was “not certain that this entity is really distinct from 


122, JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


L. bachmanii larvata (Strecker). These forms, whatever their true 
relationship, have been taken flying together (Heitzman & Heitzman, 
1972). However, W. D. Field (in litt., 11 May 1976) reports that spec- 
imens from the migration collected by Howe in Kansas are not cari- 
nenta. Examination of personal color transparencies taken during the 
1971 migration in central Texas reveal that the taxon involved was 
larvata. Ferris (1976) reports that all snout butterflies figured in Howe 
(1975) are larvata and that the verbal descriptions of these two species 
are reversed. 


1971 Observations 


As early as 20 July 1971 snout butterflies were very common in 
Austin. Butterflies were moving individually (similar to migration of 
22-25 August reported by Helfert). On 25 July snout butterflies were 
seen “by the thousands.” Many were noted on leaflets of a chinaberry 
tree (Meliaceae: Melia azedarach L.) growing on the bank of Waller 
Creek on the campus of the University of Texas at Austin. Adults 
landed on leaflets and appeared to feed from the surface as they un- 
rolled their probosces and moved them over the leaflet surface. On 
26 July numerous adults were observed in similar feeding behavior 
on leaves and stem nodes of bean plants at the Brackenridge Field 
Laboratory (BFL) of University of Texas at Austin. Possibly nutrients 
and water (Austin area was still suffering from year-long drought) 
were obtained by these butterflies. Many snout butterflies were seen 
as late as 28 July, after which time numbers of snout butterflies de- 
clined. 

A number of mature larvae of larvata were observed at BFL as 
early as 18 August on Texas sugarberry (Ulmaceae: Celtis laevigata 
Willd.) (see Neck, 1976). On 23 August adults were again noted as 
abundant (Helfert reports 22-25 August). At this point butterflies were 
traveling individually (as reported by Helfert) and were most abun- 
dantly found feeding at flowers of kidneywood (Leguminosae: Eysen- 
hardtia texana Scheele). 

An incredibly dense concentration of migrating snout butterflies 
moving NNE peaked on 27 August. Densities over central urban areas 
were not as high as observed outlying urban or rural areas. Peak num- 
bers dropped off after 30 August. Helfert reports peak on 26—28 Au- 
gust with last large numbers seen on 2 September. Helfert records no 
further observations after 2 September, but my notes record “many, 
many snout butterflies” migrating together on 20 September after an 
intervening period of low abundance. 

Indications are that this flight of snout butterflies traveled north- 


VOLUME 37, NUMBER 2 23 


TABLE 1. Drought severity in Austin, Texas, 1970-1971. 


Accumulated 


deficiency 
Recorded rainfall Normal rainfall N iene: 1970 
November 1970 ar et, ZA, 
December On DES} 4.54 
January 1971 0.04 855) 6.85 
February 0.69 2.58 8.74 
March 0.79 PAS 10.08 
April 1.07 3.55 12.56 
May om Petal 13.90 
June 1.68 S22. 15.44 
July 1523 2.18 16.39 
August 5.69 1.94 12.64 
September 213 3.44 13.95 


Total 14.80 28.75 — 


ward at least to east central Kansas, as Howe (1975:258) reported that 
“hundreds” of snout butterflies appeared in Franklin County, Kansas, 
in September and early October 1971 still “flying in a due north- 
northeast (N22.5°E) direction.” Franklin County is about 1100 km 
north of the Austin area at an approximate direction of N12°E. If one 
assumes that these were butterflies from the peak migration of late 
August-early September in central Texas, these butterflies traveled 
the distance in approximately thirty days for an average daily distance 
of about 35 km. Daylength at that season is decreasing but is about 
12 h. Allowing for low flight activity during the cool crepuscular pe- 
riod, one can assume an 8 h flight day (Helfert, 1972), yielding 4.4 
km/h. This speed is certainly exceeded by these butterflies, allowing 
for sufficient time for energy source location and utlization. Previous 
reports of speed of migrating snouts have varied from 7.2 to 24 km/h 
(Gable & Baker, 1922; Parman, 1926; Fletcher, 1926; Clench, 1965). 


Associated Weather System and Biotic Responses 


In summer 1971, central and southern areas of Texas were experi- 
encing a drought which had begun in late 1970 (see Table 1). Plant 
growth and insect populations were greatly depressed. Substantial 
rains occurred in the Border Country (Fig. 1) as early as 1 June (Table 
2). However, at this time the central Texas area was still suffering 
from drought conditions, although rain would soon begin. The entire 
July rainfall (1.23 inches vs. average 2.18) occurred on and after 24 
July. The first six days of August brought 5.23 inches of rain (average 
for entire month is only 1.94 inches). 


124 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


= alee 

= slats 

eee eae 

ater 

ry aaa ep 
bes mean are 

ee | 

vel 4 

| : 

; pobraba al 
HS | | eee nal 

Oiceccas 
Ca ie 
ore res 


y, FEO Ne - Wea 
ligase SS anal a aN ¢ 1 C Zz, 
: @SAN ANTONIO Ree 

: : aN pO Pan 

BORDER COUNTRY —-4 “< 7% nm ate 

Nis | 4 \, oes : a 
EAGLE PASS® ~— CARRIZO SPRINGS<  /! < 
Renrcne eee 


pods eee BNW A Ie 
ee Ter PET ao / 
Re 
Ree 
To i i! F 
era oa 
LB ee 


Fic. 1. Base map of Texas, showing localities discussed in text. Solid line is Bal- 
cones Escarpment Zone; arrow indicates general direction of snout butterfly migration; 
names refer to weather station locations. 


Central Texas was transformed from drought to lush conditions in 
a very short period of time. Plants put on new growth (up to 1 m 
branch growth in C. laevigata); insect populations increased dramat- 
ically. Various bird species, whose nesting had been curtailed by the 
dry spring and early summer, responded with late nesting attempts 
(Webster, 1972). Impact of the 1971 drought and heavy rain conditions 
affected the population dynamics of various lizards (Clark, 1976; Mar- 
tin, 1977), snakes (Clark, 1974; Clark & Fleet, 1976) and small mam- 
mals (Beasom & Moore, 1977). | 

Rainfall in Austin for the remainder of August totaled 0.45 inches, 
while September rainfall was sporadic and subnormal (2.13 inches vs. 
average 3.44). Therefore, plant growth occurred profusely for a brief 


VOLUME 37, NUMBER 2 125 


TABLE 2. Extensive rainfall (and associated cool temperatures) in central and south 
Texas in summer 1971 which broke drought of 1970-1971. 


Greatest 
Departure Precipi- Departure aily 


Station Ave. temp. from normal tation from normal rain total 
June 1971 
Central Texas 
Austin 83.7 +1.8 1.68 == JL 55%! 1.40 
San Antonio 83.6 +1.7 2.74 —0.21 1.07 
Border Country 
Carrizo Springs 84.2 =(0'5 13.52 + 10.98 3.81 
Del Rio 80.1 —4.3 4.87 +2.58 1.03 
Eagle Pass 82.0 —3.8 147 + 12.28 4.85 
Encinal 3NW 83.2 —].4 10.80 +8.42 4.00 
July 1971 
Central Texas 
Austin 85.9 +1.4 1.23 —0.95 0.94 
San Antonio 85.9 +1.9 1.05 —1.04 1.03 
Border Country 
Carrizo Springs 84.2 =2.4 0.17 — 1.66 0.13 
Del Rio 82.0 —4,2 0.45 —0.86 0.33 
Eagle Pass 83.0 —4.6 0.32 — 1.72 0.32 
Encinal 3NW 83.8 —2.7 0.17 —1.40 0.17 


Central Texas 


Austin 81.2 =—20 5.69 +3.75 Li 

San Antonio Shi = 25) 9.42 +7.06 2.38 
Border Country 

Carrizo Springs 80.4 =(6); Il 12.46 +10.15 6.00 

Del Rio 78.6 —7.2 6.10 +4.58 2.76 

Eagle Pass 79.2 —8.1 ANG +4.82 Wed, 

Encinal 3NW 79.8 (a5) 3.62 +1.72 0.95 


Central Texas 


Austin 79.3 =O) 2elks = IL.Sih 0.74 

San Antonio 80.1 = 5 4.57 + 1.08 1.86 
Border Country 

Carrizo Springs 80.1 —1.4 — — eit 

Del Rio one —1.4 0.50 —2.11 0.25 

Eagle Pass 78.6 NS 1.19 —1.31 0.31 

Encinal 3NW WA —3.9 9.90 +7.09 4,50 


period in early August but was terminated by a return to dry condi- 
tions. 
DISCUSSION 


Study of published reports of previous snout migrations and con- 
temporary weather systemics reveals that climatic conditions of cen- 


126 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


54 days 


Tr 


f— uaa 


AUGUST 27 
SEPTEMBER 23 
NUMBER OF BUTTERFLIES OBSERVEO(RELATIVE) 


PRECIPITATION 


JUNE JULY AUGUST SEPTEMBER OCTOBER 


Fic. 2. Time relationship between rainfall at Eagle Pass and snout butterfly migra- 
tion at Austin. Abscissa = relative butterfly numbers; Ordinate = calendar. 


tral and south Texas are the key to snout butterfly migrations (Neck, 
unpubl.). The initial low-density migration in late July (not observed 
by Helfert) probably involved adults from south Texas which devel- 
oped subsequent to the initial rains in the Border Country. Upon 
migrating to central Texas (under drought conditions) flights of these 
first generation butterflies were not joined by central Texas snout 
butterflies (which occurred in very low numbers at that time). Addi- 
tionally, stimuli which evoke massive synchronized migration were 
lacking; adults moved more or less individually. Eggs were laid by 
these south Texas butterflies on central Texas Celtis; larvae from these 
eggs faced conditions favoring rapid population growth (abundant new 
plant growth and depressed populations of natural control agents). 
Adults of this second generation formed the massive migrations seen 
in late August in Austin (note mature larvae found in mid-August at 
Austin). The smaller-scale migration seen in mid-September appar- 
ently involved adults of a third generation. The reason for its smaller 
size was probably two-fold: 1) reduced amount of new plant growth; 
and 2) increased population levels of natural control agents. 

Helfert (1972) suggests that central Texas is a major breeding ground 
of larvata. Breeding occurred in moderate numbers in 1971 in central 
Texas, but such conditions are unusual. Normally, larvata occurs at 
low population levels in central Texas. The largest breeding ground 
for this species is south Texas and adjacent parts of northeastern Mex- 
ico where its favored larval foodplant (Kendall and Glick, 1972), spiny 


VOLUME 37, NUMBER 2 127 


hackberry (Celtis pallida Ten.), is very abundant. Dorothy Yeager 
(pers. comm., 20 Feb. 1976) reported that during September and Oc- 
tober hackberry trees within a 4-km radius of Pearsall, Frio County, 
were “almost completely denuded of leaves.” This observation also 
indicates that the Border Country area is the breeding grounds for a 
great number of the snout butterflies of this migration. 

Occurrence of associated major and minor migrations of the snout 
butterfly has apparently not been reported previously. Temporal sep- 
aration of occurrence of drought-breaking rains in south and central 
Texas may have caused the occurrence of these major and minor mi- 
grations. Normally, such drought-breaking rains occur in these areas 
concurrently. The temporal relationship between the drought-break- 
ing rains and the snout butterfly migrations is shown in Fig. 2. 


QUESTIONS RAISED 


A series of questions has been raised by analysis of this snout but- 
terfly migration. Tentative answers are here given to these questions. 
Study of future migrations will shed light on the validity of these 
answers. 

1) Would the late July (first minor) migration have been a larger 
migration if rains had occurred in central Texas in late June? 

Not likely; two generations of favorable breeding conditions are 
probably required to produce such a tremendous build-up of numbers 
of individuals. The central Texas area was not the source of these 
butterflies except for a contribution of unknown importance to the 
late August (major) migration. 

2) Would the migration of late August (major) have occurred if rains 
had not returned to the Border Country area in late July and early 
August? 

Yes, although possibly on a smaller scale; these butterflies origi- 
nated as the second generation from the Border Country of Texas and 
Mexico as a result of the June rains. Rainfall in late July in the Border 
Country was relatively minor. One possible result of the absence of 
these later rains would have been the diminution of this major mi- 
gration. 

3) Would the late September (second minor) migration have been 
a major migration if rain had continued through September? 

Probably not; by this time natural control agents had presumably 
increased in population size sufficiently to have a dampening effect 
upon population levels of the snout butterfly. One should also con- 
sider the possibility that many of these butterflies may have originated 
in central Texas. 


128 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


ACKNOWLEDGMENTS 


For information incorporated into this report, I thank W. D. Field, W. H. Howe and 
D. Yeager. T. D. Samsell, III drafted Fig. 2. 


LITERATURE CITED 


BEASOM, S. L. & R. A. MOORE. 1977. Bobcat food habit response to a change in prey 
abundance. Southwestern Nat. 21:451-454. 

CLARK, D. R., JR. 1974. The western ribbon snake (Thamnophis proximus): ecology 
of a Texas population. Herpetologica 30:372-379. 

1976. Ecological observations on a Texas population of six-lined racerunners, 
Cnemidophorus sexlineatus (Reptilia, Lacertilia, Teiidae). J. Herpetology 10:133- 
138. 

CLARK, D. R., JR. & R. R. FLEET. 1976. The rough earth snake (Virginia striatula): 
ecology of a Texas population. Southwestern Naturalist 20:467-478. 

CLENCH, H. K. 1965. A migration of Libytheana and Kricogonia in southern Texas. 
J. Lepid. Soc. 19:223-224. 

FERRIS, C. D. 1976. The butterflies of North America (book review) J. Lepid. Soc. 
30: 138-143. 

FLETCHER, R. K. 1926. Notes on a migration of the snout butterfly (Lepid., Liby- 
theidae). Entomol. News 37:106—107. 

GABLE, C. H. & W. A. BAKER. 1922. Notes on a migration of Libytheana bachmanii 
Kirtl. Can. Entomol. 54:265-—266. 

HELFERT, M. R. 1972. Migratory behavior of the snout butterfly, Libytheana bach- 
manii larvata. (Strecker). Entomol. News 83:49-52. 

Howe, W. H. 1972. Family Libytheidae. The snout butterflies. Pp. 257-258, in W. 
H. Howe (coord. ed. and illus.). The Butterflies of North America. Doubleday & 
Co., Garden City, New York. 

KENDALL, R. O. & P. A. GLick. 1972. Rhapalocera collected at night in Texas. J. Res. 
Lepid. 10:273-283. 

KENDALL, R. O. & C. A. KENDALL. 1971. Lepidoptera in the unpublished field notes 
of Howard George Lacey, naturalist (1856-1929). J. Lepid. Soc. 25:29-44. 

MARTIN, R. F. 1977. Variation in reproductive productivity of range margin tree 
lizards (Urosaurus ornatus). Copeia 1977:83-92. 

NECK, R. W. 1976. Lepidopteran foodplant records from Texas. J. Res. Lepid. 15:75— 
82. 

PARMAN, D.C. 1926. Migrations of the long-beaked butterfly, Libythea bachmanii 
Kirtland (Lepid.: Libytheidae). Entomol. News 37:101—-106. 

WEBSTER, F. S., JR. 1972. Fall migration 1971. South Texas Region Amer. Birds 26: 
84-88. 


Journal of the Lepidopterists’ Society 
37(2), 1983, 129-139 


LEPIDOPTERA ASSOCIATED WITH WESTERN SPRUCE 
BUDWORM IN THE SOUTHWESTERN UNITED STATES 


ROBERT E. STEVENS 


Rocky Mountain Forest and Range Experiment Station, USDA Forest Service, 
240 W. Prospect St., Fort Collins, Colorado 80526! 


V. M. CAROLIN 
9030 S.E. Mill Street, Portland, Oregon 97216 


CATHERINE STEIN 


Southwestern Region, USDA Forest Service, Federal Building, 
517 Gold Ave. S.W., Albuquerque, New Mexico 87102 


ABSTRACT. Western spruce budworm, Choristoneura occidentalis Freeman (Tor- 
tricidae), is an important pest of Douglas-fir and white fir in the southwestern United 
States. A variety of other Lepidoptera, several previously unrecognized from this part 
of the country, commonly occupy similar feeding niches as larvae. Included are species 
of Geometridae, Gelechiidae, Noctuidae, Plutellidae, Pyralidae, and Tortricidae. Notes 
are presented on species life history, and field identifying features in late larval and 
adult stages. 


Western spruce budworm, Choristoneura occidentalis Freeman, is 
an important defoliator of Douglas-fir, Pseudotsuga menziesii (Mirb.) 
Franco, and true firs, Abies spp., throughout western North America. 
It also feeds on spruces, Picea spp., and western larch, Larix occi- 
dentalis Nutt. (Furniss & Carolin, 1977). The budworm is a common 
pest of Douglas-fir and white fir, A. concolor (Gord. & Glend.) Lindl. 
ex Hildebr., and is currently in outbreak status in northern Arizona 
(Kaibab Plateau, Coconino County), in northern New Mexico (Jemez 
and Sangre de Cristo Mountains, Sandoval and Taos Counties, re- 
spectively), and in Colorado (mainly in the Front Range of the Rocky 
Mountains—Larimer, Boulder, Jefferson, Teller, Fremont and Custer 
Counties). 

A variety of other Lepidoptera coexist with the budworm, occupy- 
ing similar feeding niches in the larval stage. These associates have 
been little known in this part of the country. The main objective here 
is to summarize this information for other workers, so that with al- 
ready available keys to larvae (Carolin & Stevens, 1979, 1981), they 
can identify common budwormm associates and have information about 
each species life history and habits. 

Other Lepidoptera may sometimes occur in sufficient numbers to 
also qualify as budworm associates. However, the ones discussed here 
are present more or less regularly, and are considered to be the 


‘ Headquarters is in Fort Collins, in cooperation with Colorado State University. 


130 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


common set of associates in the area. Table 1 lists these species. All 
probably have the ability to colonize both main budworm hosts, Abies 
and Pseudotsuga. | 

Larvae of other insect groups, including Xyelidae, Diprionidae, and 
Pamphiliidae (Hymenoptera), are sometimes found feeding on foliage 
along with budworms. These are readily separable from Lepidoptera 
larvae, however, and are not considered here. Also excluded is the 
Douglas-fir tussock moth, Orgyia pseudotsugata (McDunnough); al- 
though sometimes common on the same hosts, the tussock moth oc- 
cupies a different feeding niche and is not usually a budworm asso- 
ciate. 

Some of the species discussed here are well-known forest insects 
and have been studied elsewhere in North America. For these, per- 
tinent information is summarized to help in field identification and 
an understanding of life histories as the insects relate to western spruce 
budworm. In several cases little is known about the species’ life his- 
tory and habits, and in some of these new information is presented. 
Incorporated also are pertinent and previously unpublished obser- 
vations made by Carolin in the Pacific Northwest. 

Voucher specimens are kept in the insect museum at the Rocky 
Mountain Forest and Range Experiment Station, Fort Collins, Colo- 
rado. 


Gelechiidae 
Chionodes abella (Bsk.) 


C. abella (Fig. la) is a rare species reared from white fir in the 
Jemez Mountains. A small (wingspan 15 mm), strikingly-patterned 
moth, C. abella is not likely to be confused with any of the other 
common budworm associates. The larva is mostly greenish-brown, 
with a tan head capsule. The thoracic legs and posterior part of the 
first thoracic segment are black. Details of its life history and habits 
are unknown. 


Coleotechnites sp. 


The genus Coleotechnites includes several well-known forest pests 
(e.g., the needle miners C. milleri (Busck) and C. starki (Freeman)), 
(Furniss & Carolin, 1978), as well as several undescribed species (R. 
W. Hodges, pers. comm., 1981). One or more of the latter are bud- 
worm associates, found extensively throughout the western United 
States. The moths (Fig. lb) are small (wingspan 10-12 mm), and most- 
ly black and white. We have a few specimens from the Jemez Moun- 
tains. 


VOLUME 37, NUMBER 2 131 


TABLE 1. Lepidoptera associated with western spruce budworm in the southwest- 
ern United States. 


Family Species 
Gelechiidae Chionodes abella (Busck) 
Coleotechnites sp. 
Geometridae Enypia griseata Grossbeck 
Eupithecia catalinata McDunnough 
Noctuidae Achytonix epipaschia (Grote) 


Syngrapha angulidens Smith 
Egira (=Xylomyges) simplex (Walker) 


Plutellidae Ypsolophus nella (Busck) 
Pyralidae Dioryctria spp. 
Tortricidae Acleris gloverana (Walsingham) 


Argyrotaenia dorsalana (Dyar) 
Argyrotaenia klotsi Obraztsov 
Argyrotaenia provana Kearfott 

Clepsis persicana (Fitch) 

Griselda radicana (Heinrich) 

Zeiraphera hesperiana Mutuura & Freeman 


Geometridae 


Enypia spp. 

Although they never appear to occur in large numbers, loopers of 
the genus Enypia are widely distributed budworm associates on both 
Abies and Pseudotsuga. Evans (1960) indicates that E. griseata Gross- 
beck and E. venata (Grote) are found in the Southwest; we have 
occasionally reared griseata. Adults of both species are large (wing- 
span 35-39 mm) gray moths, and are difficult to tell apart by non- 
specialists. E. griseata is shown in Fig. lc. Eggs of both species are 
ivory colored when first laid. They are laid on needles, singly or oc- 
casionally in pairs. According to Evans (1960), the larvae are solitary 
and constitute the overwintering stage, and fully-developed larvae of 
the two species differ as follows: 


E. griseata E. venata 
Head pale green-brown. Head brown, irregularly patterned. 
Body green dorsally; venter paler green. | Body pale golden brown dorsally; over- 
Narrow pale-green dorsal line; wide all pattern of broken dark irregular 
near-white sub-dorsal stripes. lines; posterior parts of segments 


darker than anterior portions, darker 
reddish-brown irregular dorsal stripe. 


Pupation is on the foliage, in a loosely-constructed cocoon. 


JOURNAL OF THE.LEPIDOPTERISTS’ SOCIETY 


Fic. 1. Budworm associate adults: a, Chionodes abella; b, Coleotechnites sp.; e, 
Enypia griseata; d, Eupithecia catalinata; e, Achytonix epipaschia; f, Syngrapha 
angulidens; g, Egira simplex; h, Dioryctria sp.; i, j, Ypsolophus nella; k, Argyro- 


VOLUME 37, NUMBER 2 


taenia dorsalana; 1, A. klotsi; m, A. provana; n, Clepsis persicana; o, Griselda radi- 
cana; p, Zeiraphera hesperiana. 


134 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Eupithecia catalinata McD. 


E. catalinata is a little-known species of looper not previously rec- 
ognized as a budworm associate. However, many members of the 
genus feed on coniferous foliage (McGuffin, 1958), and one, E. an- 
nulata (Hulst), is a recognized budworm associate on the West Coast 
(Carolin, 1980). The adults (Fig. 1d) are small (wingspan 20-22 mm) 
gray moths with indistinctly marked wings. 


Noctuidae 
Achytonix epipaschia (Grote) 


A. epipaschia has been reared in small numbers from the Jemez 
Mountains. Carolin (1980) considers it a “sporadic” and “occasional” 
budworm associate in the Pacific Northwest. The distinctively marked 
gray moths (Fig. le) have a wingspan of 25-30 mm. They fly about 
the same time as does the budworm, and the two species have some- 
what similar life histories; both overwinter as small larvae and pupate 
on the shoots where they have been feeding. Early instars of Achy- 
tonix also resemble those of budworm. However, in late instars, the 
green abdomen, conspicuous black setal bases, and three broad lon- 
gitudinal lines on the dorsum make Achytonix readily identifiable. 


Syngrapha angulidens (Smith) 


S. angulidens (Fig. 1f), a large (32-34 mm wingspan), distinctively- 
marked noctuid, is another relatively uncommon budworm associate 
in the Southwest. Little is known of its life history and habits, but 
Eichlin & Cunningham (1978) indicate that eggs are deposited singly 
and larvae overwinter. Larvae of Noctuidae possess varying numbers 
(3-5 pairs) of abdominal prolegs. Some with three pairs move like 
geometrids; S. angulidens is one of these. 

Egira simplex (Wlk.) 

E. simplex is widely distributed throughout the West (Furniss & 
Carolin, 1977) and is an occasional budworm associate in the South- 
west. Its life history is much like that of the budworm, except that 
E. simplex overwinters as a pupa in the soil. The adult (Fig. 1g) is a 
large (38-40 mm wingspan) gray moth. Fully developed larvae are 


up to 35 mm long, green with white longitudinal dorsal and sub- 
dorsal lines, and shiny black head capsules and dorsal and anal 


shields. 


Plutellidae 
Ypsolophus nella (Bsk.) 


The life history and habits of Y. (=Abebaea) nella have not previ- 
ously been described. However, records on file in the insect museum 


VOLUME 37, NUMBER 2 135 


Fic. 2. White fir needles tied into “tents” by larvae of Ypsolophus nella. 


at the Rocky Mountain Forest and Range Experiment Station indicate 
that the species is common on Abies concolor throughout Colorado 
and New Mexico. It has been a consistent budworm associate on white 
fir in our recent collections. Carolin (1980) reported Y. prob. cervella 
(Walsingham) as a rarely collected associate on Douglas-fir in western 
Oregon in the 1950’s. 

Adults and larvae of Y. nella are highly distinctive. The adult (Figs. 
li, j) is a small (wingspan 20-2] mm) moth, with gray abdomen and 
hindwings, and narrow brown forewings ornamented by longitudinal 
lines made up of black and white scales. The amount of black on the 
wings varies and may be totally lacking in some individuals. 

Fully developed larvae, about 15 mm long, are generally purplish 
to pale green, with two narrow and one broad yellowish-green lon- 
gitudinal lines on each side of the dorsal midline. Black setae and 


136 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


setal bases also constitute distinctive recognition characters. The lar- 
vae are particularly active and capable of unusually rapid movement 
when disturbed. 

The character of larval feeding is also distinctive. The ends of the 
needles are webbed together soon after their emergence from the 
bud. As the needles elongate, their central parts diverge, creating an 
expanding “tent” (Fig. 2) within which the larva feeds. Pupation oc- 
curs in the foliage in a loosely constructed cocoon. Summer larval and 
pupal periods approximate that of the budworm; eggs have not been 
seen nor is the overwintering stage known. 


Pyralidae 
Dioryctria spp. 


The spruce coneworm, D. reniculelloides Mutuura & Munroe, has 
long been recognized as a budworm associate, sometimes occurring 
in great numbers. Carolin (1980) reported as many as 158 Dioryctria 
larvae per 100 buds in a 1957 sample plot in central Oregon. Both it 
and insects identified as D. pseudotsugella Munroe are commonly 
reared along with budworms in the Southwest. The two Dioryctria 
species are difficult for the non-specialist to separate either as larvae 
or adults; for the purposes of this article Dioryctria associates are 
conatlered a single entity. 

Moths have gray forewings with distinctive transverse bands (Fig. 
lh), and are not likely to be confused with any of the other budworm 
associates. Although we have reared adults with wingspans as small 
as about 15 mm, most specimens are larger, 20-25 mm. Larvae are 
also distinctive; the dorsum of well-developed individuals is gener- 
ally pinkish to reddish-brown, with broad, irregular white and black 
lines on either side of the dorsal midline. The pupa is dark brown to 
black and is found in the foliage. In general, the life history parallels 
that of the budworm; however, Dioryctria may pupate slightly later. 


Tortricidae 
Acleris gloverana (Wlshm.) 


A. gloverana, the western blackheaded budworm, has not previ- 
ously been known from the southwestern United States; however, it 
has been fairly common in Jemez Mountains rearings of budworm 
associates. A. gloverana is a serious forest pest in British Columbia 
and southeast Alaska (Furniss & Carolin, 1977), and its life history 
and habits have been thoroughly studied in that region. Also, Powell 
(1962) provides a detailed discussion of it. 

Adults, wingspan 18-22 mm, display a bewildering variety of fore- 


VOLUME 37, NUMBER 2 Sh 


wing markings, making identification difficult for the inexperienced 
observer. Furniss & Carolin (1977) show three of the more common 
morphs. In general, the moth is dark colored; the forewings are var- 
iously marked with brown, white, yellow, and orange. Small larvae 
have black head capsules and prothoracic shields, and lemon-yellow 
bodies. The latter instars have chestnut-brown head capsules and grass- 
green bodies. 

The life cycle and habits are similar to those of the western spruce 
budworm; however, A. gloverana eggs are laid singly on needles, and 
the egg overwinters. On the West Coast, Acleris adults emerge 2-3 
weeks later than spruce budworms. 


Argyrotaenia spp. 


According to Hodges (in litt.), the genus Argyrotaenia includes 34 
North American species. Of these, A. dorsalana (Dyar), A. klotsi Obr., 
and A. provana Kearf. are budworm associates. All are found regularly 
in the Southwest. These species are sufficiently similar in most re- 
spects to justify treating them together. 

Except for A. dorsalana, details of their life histories are essentially 
unknown; however, they are probably all similar. Eggs of A. dorsa- 
lana are laid in overlapping rows on needles, much as in the case of 
the budworm. The eggs are slightly smaller and are finer-textured 
than those of budworm, and the egg mass usually has an orange-pink 
tint. The small larva overwinters. Larval feeding is also similar to that 
of budworm; pupation is in the foliage, slightly earlier than budworm. 

Larvae of all species are generally green. The moths, while differ- 
ently marked, are all about the same size, wingspan 20-25 mm. A. 
dorsalana is generally the most common member of the genus as a 
budworm associate in the Southwest. Forewings of the adult (Fig. 1k) 
are largely straw-yellow but exhibit a variety of brown markings. The 
most common form is nearly pure yellow with a small marking on the 
posterior margin. Some have no marks at all; others are heavily pat- 
terned. 

The forewings of A. klotsi and A. provana (Figs. 11, m) are gray- 
black, with distinctive white (provana) or yellow (klotsi) bands and 
patches. Adults of these species appear to show much less morpho- 
logical variation than do those of A. dorsalana. 


Clepsis persicana (Fitch) 


C. persicana (Fig. In) is a striking species we reared only once from 
the Jemez Mountains. The forewings (wingspan 18 mm) of the adult 
are orange to ochreous-orange basally, having a dark gray “V’’-shaped 
section distally and white patches on the anterior margin and apex of 


138 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


the wingtip. The hindwings are gray dorsally and white ventrally. 
Fully developed larvae are about 12-15 mm long and generally green; 
the head capsule is green with a brownish tint; prothoracic and anal 
shields are emerald-green; the dorsum is dark olive-green with two 
whitish longitudinal lines and whitish setal areas. The venter is lighter 
green. Feeding habits are similar to that of budworm. 

Powell (1964) indicates that C. persicana utilizes a variety of food 
plants other than conifers; however, he also has more recently reared 
it as a budworm associate in California (J. A. Powell, unpublished 
data). Carolin reared it once from the Blue Mountains in northeastern 
Oregon and several times from Abies balsamea L. in Maine. 


Griselda radicana (Heinr.) 


G. radicana, the spruce tip moth, is another common budworm 
associate not previously known to occur in the Southwest. We reared 
several specimens from the Jemez Mountains, and presumably the 
species occurs much more generally. G. radicana is a small moth, 
wingspan 12-16 mm, having gray forewings with distinctive rusty- 
colored basal sections (Fig. lo). Young larvae are pale yellow overall. 
Later instars have the dorsum marked with three orange-brown to 
orange-red lines; fully developed larvae undergo a quiescent prepu- 
pal period, during which the abdomen becomes whitish and the lines 
disappear. Adult emergence is in late summer. Eggs, laid singly at 
the bases of needles, overwinter. Larval feeding is similar to that of 
budworm. 


Zeiraphera hesperiana M. & F. 


Z. hesperiana, commonly known as the Douglas-fir bud moth, is 
well known as a budworm associate. However, it has not previously 
been reported from the Southwest, and the only published informa- 
tion on its life history is a brief mention by Carolin (1980). Mutuura 
& Freeman (1966) described the species from British Columbia; Fur- 
niss & Carolin (1977) also record it from Oregon. We have specimens 
from Idaho and Montana, and from the Jemez and Sangre de Cristo 
ranges in New Mexico. Thus the species appears to have a wide dis- 
tribution. The following notes on life history and habits are largely 
from Carolin’s observations in Oregon. Stein and Stevens have noted 
similar habits in New Mexico. 

The adult (Fig. lp) is a distinctly marked, generally dark moth, 
wingspan 15-20 mm. At rest it is readily separable from moths of other 
common budworm associates by the presence of a prominent saddle- 
like white to brownish-white patch located centrally on the forewings. 
The forewings are otherwise marked with characteristic patches made 


VOLUME 37, NUMBER 2 139 


up of black, cream, brown, and orange-brown scales. Eggs overwinter. 
They are yellow, spiny, and laid singly on bark of limbs, 50 cm or 
more back from branch tips. In spring, new larvae enter buds and 
feed therein, concealed until the buds open. Feeding becomes visible 
as shoots develop. Fully developed larvae are 12-15 mm long and 
generally stout in form. The head capsule and prothoracic shield are 
golden to chestnut-brown; the abdomen is generally yellowish, with 
a broad, olive-brown to chocolate-brown dorsal stripe. The prothora- 
cic shield usually has a characteristic black posterior margin. Larvae 
leave the feeding area to pupate in the soil or duff layer, well before 
the time of budworm pupation. 


ACKNOWLEDGMENTS 


The authors thank Pamela Farrar for assistance in the field in 1979, and S. A. Mata 
for taking the photos. J. A. Powell, University of California, Berkeley, T. D. Eichlin, 
California Department of Food and Agriculture, Sacramento, D. C. Ferguson and R. 
W. Hodges, Systematic Entomology Laboratory, USDA, Washington, D.C., and A. Mu- 
tuura, Canada Department of Agriculture, Ottawa, kindly identified specimens. W. E. 
Waters, and W. J. A. Volney and associates, University of California, Berkeley, and M. 
J. Stelzer, Pacific Northwest Forest Experiment Station, Corvallis, Oregon provided 
very helpful study material from their rearings in New Mexico and Arizona, respec- 
tively. The material reported here was developed with partial support from the Can- 
ada—United States Spruce Budworms Program (CANUSA). 


LITERATURE CITED 


CAROLIN, V. M. 1980. Larval densities and trends of insect species associated with 
spruce budworms in buds and shoots in Oregon and Washington. USDA For. Serv. 
Res. Pap. PNW-273. Pac. Northwest For. and Range Expt. Stn., Portland, Oregon. 
18 pp. 

CAROLIN, V. M., JR. & R. E. STEVENS. 1979. Key to small lepidopterous larvae in 
opening buds and new shoots of Douglas-fir and true firs. USDA For. Serv. Res. 
Note RM-365. Rocky Mtn. For. and Range Expt. Stn., Ft. Collins, Colorado. 4 pp. 

1981. Key to large lepidopterous larvae on new foliage of Douglas-fir and true 
firs. USDA For. Serv. Res. Note RM-365. Rocky Mtn. For. and Range Expt. Stn., 
Ft. Collins, Colorado. 4 pp. 

EICHLIN, T. D. & H. B. CUNNINGHAM. 1978. The Plusiinae (Lepidoptera: Noctuidae) 
of America north of Mexico, emphasizing genitalic and larval morphology. USDA 
Tech. Bull. 1567. 122 pp. 

EVANS, DAviIpD. 1960. A revision of the genus Enypia (Lepidoptera: Geometridae). 
Ann. Entomol. Soc. Am. 53:560-574. 

FuRNISS, R. L. AND V. M. CAROLIN. 1977. Western forest insects. USDA For. Serv. 
Misc. Publ. 1339. Washington, D.C. 654 pp. 

MCGUFFIN, W. C. 1958. Larvae of the nearctic Larentiinae (Lepidoptera: Geometri- 
dae). Canad. Entomol. 90, Suppl. 8. 104 pp. 

MutTuura, A. & T. N. FREEMAN. 1966. The North American species of the genus 
Zeiraphera Treitschke (Olethreutidae). J. Res. Lepid. 53:153-176. 

POWELL, J. A. 1962. Taxonomic studies on the Acleris gloverana-variana complex, 
the black-headed budworms (Lepidoptera: Tortricidae). Canad. Entomol. 94:833- 
840. 


1964. Biological and taxonomic studies on tortricine moths, with reference to 
the species in California. Univ. of Calif. Pubs. Entomol., Vol. 32. 317 pp. 


Journal of the Lepidopterists’ Society 
37(2), 1983, 140-145 


TWO NEW SPECIES OF THE TRIBE EUCOSMINI 
(TORTRICIDAE) CLOSELY RELATED TO 
PHANETA GRANULATANA (KEARFOTT) 


ANDRE BLANCHARD 
3023 Underwood, Houston, Texas 77025 


AND 


EDWARD C. KNUDSON 
804 Woodstock, Bellaire, Texas 77401 


ABSTRACT. Phaneta linitipunctana and Phaneta argutipunctana are described 
and imagines and male and female genitalia are figured. Phaneta granulatana imag- 
ines and genitalia are also figured. 


Phaneta linitipunctana, new species 


(Figs. 1-7) 


Head. Front and vertex light ochreous. Labial palpi exceeding front by 24% eye 
diameters, light ochreous, with brushlike 2nd segment obscuring blackish 3rd segment. 
Antennae simple, light ochreous. 

Thorax. Light ochreous with fulvous spots on patagia and mesonotum. 

Forewing (Figs. 1-4). Ground color light ochreous with extensive fulvous macula- 
tion, which, in well marked examples shows a tendency to form vertical rows. On the 
basal third, extending from dorsum to % the distance to costa, the fulvous markings are 
heavier, forming an ill-defined patch, the outer margin of which is angled slightly 
outward from dorsum. A narrow streak of ground color extends along the fold, inter- 
rupting the fulvous maculation. Ocelloid patch with central area lighter than ground, 
with a pearly luster, and bearing a weak scattering of black scales, tending to form 
three horizontal dashes. Along upper margin of ocelloid patch is a small elongate patch 
composed of small flat, white tipped brown scales. Fringe consists of two bands, the 
inner band fairly broad and composed of scales having a whitish base and tip with a 
dark brown center; the outer band is ochreous. 

Hindwing. Light fuscous. Fringe with fuscous inner band and ochreous outer band. 

Length of forewing. Males (n = 12): 6.6-8.3 mm, average 7.3 mm. Females (n = 9): 
7.3-8.7 mm, average 8.0 mm. 

Venation. Hindwing: M3 and Cul stalked for % the length of Cul. Rs and M1 
approximate for % the length of M1. 

Male genitalia. As in Figs. 5 and 6. 

Female genitalia. As in Fig. 7. 

Holotype. Male, Nueces Co., Texas, North Padre Island, 9-IX-74, slide A.B. 4342, 
collected by A. & M. E. Blanchard and deposited in the National Museum of Natural 
History. 

Paratypes. Hemphill Co., Texas, Canadian, 13-VIII-71, 2 males (slides A.B. 2905, 
2844), 2 females (slide A.B. 4999), 15-VIII-71, 3 females (slides A.B. 2845, 4344, 5053); 
Nueces Co., Texas, North Padre Island, 9-IX-74, 3 males (slide A.B. 4343), 29-IX-75, 1 
male, 17-VIII-76, 1 male (slide A.B. 4306), 24-IX-79, 2 males (slide A.B. 5028), all 
collected by A. & M. E. Blanchard; Nueces Co., Texas, North Padre Island, 12-X-79, 
2 males (slides ECK 197,199), 1 female; Cameron Co., Texas, 10 miles west of Browns- 
ville, 13-X-79, 1 female, all collected by E. C. Knudson. 


VOLUME 37, NUMBER 2 14] 


Fics. 1-7. Phaneta linitipunctana: 1, paratype male, North Padre Island, Nueces 
Co., Texas, 9-IX-74; 2, holotype male, same data, slide A.B. 4342; 3, paratype female, 
Canadian, Hemphill Co., Texas, 13-VIII-71; 4, paratype female, same locality, 15-VIII- 
71, slide A.B. 4344 (all four adults same scale); 5, male genitalia of holotype, slide 
A.B. 4342; 6, male genitalia of paratype, slide A.B. 4306, North Padre Island, Nueces 
Co., Texas, 17-VIII-76; 7, female genitalia of paratype, slide A.B. 5053, Canadian, 
Hemphill Co., Texas, 15-VIII-71. The segments in Figs. 5-7 equal 1 mm. 


142 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Fics. 8-14. Phaneta argutipunctana: 8, holotype male, Canadian, Hemphill Co., 
Texas, 15-VIII-71, slide A.B. 2846; 9, paratype male, Padre Island Nat. Seashore, Kle- 
berg Co., Texas, 13-X-79; 10, paratype female, same data as holotype, slide A.B. 5000; 
11, paratype female, Camp Strake, Montgomery Co., Texas, 14-IX-77, slide A.B. 4997 
(all four adults same scale); 12, male genitalia of holotype, slide A.B. 2846; 13, male 
genitalia of paratype, slide ECK 198, North Padre Island, Nueces Co., Texas, 12-X-79; 
14.,, female genitalia of paratype, slide A.B. 4997, same data as Fig. 11. The segments 
in Figs. 12-14 equal 1 mm. 


VOLUME 37, NUMBER 2 143 


Fics. 15-19. Phaneta granulatana: 15, cotype male, Oslar, Denver, Colo., slide 
USNM 25200; 16, cotype female, Colo., 2298, slide USNM 25201; 17, male genitalia 
of cotype, slide USNM 25200, same data as Fig. 15; 18, male genitalia of cotype, slide 
USNM 25202, Oslar, Denver, Colo.; 19, female genitalia of cotype, slide USNM 25203, 
Oslar, Platte Canon, Colo. The segments in Figs. 17-19 equal 1 mm. 


Phaneta argutipunctana, new species 


(Figs. 8-14) 


Head. Front and vertex whitish with some brownish tipped scales on vertex. Labial 
palpi exceeding front by three eye diameters, whitish ochreous with some grayish 
scales on 2nd segment. 3rd segment blackish, usually hidden by 2nd. Antennae simple, 
whitish, with prominent black scaling on inner surface of scape. 

Thorax. Patagia and mesonotum whitish with dark brown central patches. 

Forewing (Figs. 8-11). Ground color whitish with pearly luster. Maculation consists 
of strongly contrasted black to dark brown scales generally arranged in evenly spaced, 
interrupted, vertical rows, except on outer third, where there is a tendency to form 
longitudinal rows which extend basad from the upper and lower outer margins of the 
ocelloid patch. Ocelloid patch ochreous with three weak black dashes near center. 
Along inner margin of ocelloid patch is a short vertical black line. Above ocelloid patch 


144 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


and indenting it along the lower portion of the outer margin are patches of small, flat, 
white tipped brown scales. Fringe consists of two bands; the inner band composed of 
dark brown scales with whitish base and tips, the outer band ochreous. The outer band 
stops short of the extreme apex, where it is replaced by the bicolored scales of the 
inner band. Termen is slightly concave. 

Hindwing. Light fuscous. Fringe darker fuscous inwardly, whitish outwardly. 

Length of forewing. Males (n = 18): 4.7-6.4 mm, average 5.7 mm. Females (n = 
14): 5.2-7.0 mm, average 6.1 mm. 

Venation. Hindwing: M3 and Cul stalked for 4% the length of Cul. Rs and M1 
approximate for %4 the length of M1. 

Male genitalia. As in Figs. 12 and 13. 

Female genitalia. As in Fig. 14. 

Holotype. Male, Hemphill Co., Texas, Canadian, 15-VIII-71, slide A.B. 2846, col- 
lected by A. & M. E. Blanchard and deposited in the National Museum of Natural 
History. 

Paratypes. Hemphill Co., Texas, Canadian, 15-VIII-71, 2 females (slides A.B. 4998, 
5000); Nueces Co., Texas, North Padre Island, 30-IX-75, 2 males; Kleberg Co., Texas, 
Padre Island National Seashore, 13-X-79, 3 males, 2 females (slide A.B. 4996); Mont- 
gomery Co., Texas, Camp Strake, Conroe, 7-IX-77, 1 male, 1 female, 14-X-77, 5 males 
(slides A.B. 4257, 5026), 3 females (slide A.B. 4997); Anderson Co., Texas, Tennessee 
Colony, 27-VIII-78, 1 male (slide A.B. 5027), all collected by A. & M. E. Blanchard; 
Nueces Co., Texas, North Padre Island, 1-X-77, 1 female, 12-X-79, 6 males (slide ECK 
198), 4 females (slides ECK 196, 200); Cameron Co., Texas, Laguna Atascosa NWR, 
13-X-79, 1 female, all collected by E. C. Knudson. 


Discussion 


These two new species are extremely close to Phaneta granulatana 
(Kearfott) (Figs. 15-19) but are separable by characteristics of the 
imagines and male genitalia. P. argutipunctana can be easily diag- 
nosed by the presence of black scaling on the scape, lacking in the 
other two species. In granulatana the ground color of the forewing 
is whitish yellow with maculation consisting of patches of dark brown 
scales, which tend to be more irregular and not arranged in vertical 
rows as in argutipunctana. The heavier maculation on the basal third 
of the forewing of linitipunctana, which tends to form an angulate 
basal patch, is lacking in the other two species. In granulatana the 
ocelloid patch is poorly defined, due mainly to the absence of the 
patch of small, flat, white tipped, brown scales above the ocelloid 
patch, which is found in the other two species. P. granulatana and li- 
nitipunctana are the same in average length of forewing; whereas, ar- 
gutipunctana is significantly smaller. The male genitalia of all three 
species are very similar, but in argutipunctana the neck of the valva 
is broader with a shallower ventral excavation. In granulatana the 
uncus tends to be narrower and more compact than in the other two 
species. 


ACKNOWLEDGMENTS 


The authors are extremely grateful to Dr. J. F. Gates Clarke of the NMNH for ar- 
ranging the loan of type specimens and for examining the manuscript and type series. 


VOLUME 37, NUMBER 2 145 


We also wish to thank the National Park Service and the Texas Parks and Wildlife 
Department for their continued assistance and cooperation. 


LITERATURE CITED 


HEINRICH, C. 1923. Revision of the North American moths of the subfamily Eucos- 
minae of the family Olethreutidae. USNM Bulletin 123:1-286, pls. 1-59. 


Journal of the Lepidopterists’ Society 
37(2), 1983, 145 


GENERAL NOTE 


OVERWINTERING AGGREGATIONS OF HACKBERRY CATERPILLARS 
(ASTEROCAMPA CLYTON: NYMPHALIDAE) 


Hackberry caterpillars (Asterocampa spp. Roeber: Nymphalidae) overwinter as mid- 
instars, presumably in fallen leaves and crevices of bark (Scudder, 1893, Guide to the 
Commoner Butterflies, Holt, New York). In the fall of 1981 in Gainesville (Alachua 
Co.), Florida, the preparatory overwintering behavior of Asterocampa clyton Boisduval 
and Leconte was observed. After molting in mid- to late October, the greenish cater- 
pillars moved up to several meters from the molting site to the ends of branches of 
their host plant, hackberry (Celtis laevigata Willdenow). Each group of caterpillars 
effectively tied a leaf to its branch by repeatedly laying silk over the junction of the 
branch and leaf petiole. Some groups tied the sides of the leaf together. Eventually 
the leaf curled and dried around the caterpillars. By late fall most of the leaves still on 
the trees were those tied by the caterpillars. Occasionally, the aggregation of caterpil- 
lars split and, consequently, two or more leaves at the end of a branch were tied, each 
leaf sheltering some caterpillars. By this time the caterpillars were pinkish-brown, 
blending with the dead leaves. 

To determine the mean number of larvae per overwintering group, 20 groups were 
collected in December (just after leaf abscission) and 21 groups in late February (just 
prior to budbreak). Group size was not significantly different (x = 8.7 larvae + 1.9 S.E. 
in December and x = 10.1 + 2.7 S.E. in February; Mann-Whitney U test, P > 0.20). 
Similar group size early and late in the overwintering period suggests that probably 
few individuals were lost from an aggregation during that period. 

To determine the effectiveness of tying leaves to the trees for overwintering sites, 
larval groups were marked in December by attaching numbered, plastic bird bands to 
the branches. Of 71 groups, 16% were recovered in late February, each with more than 
half of the leaf and caterpillars present. Fourteen percent of the markers had less than 
half of a leaf and 70% of the markers had no leaf. None of these had caterpillars. This 
supports the idea that disappearance from the branches was a larval group event rather 
than an individual event. 

Thus, it appears that hackberry caterpillars overwintered within leaves tied to branch 
tips on their host plants when more than half of the leaf remained intact and tied to 
the branch. Tied leaves and aggregations of caterpillars missing from the trees may be 
a result of avian predation or weather, causing deterioration or detachment of the leaves. 


NANcy E. STAMP, Department of Zoology, University of Florida, Gainesville, Flor- 
ida 32611. (Present address: Department of Zoology, University of California, Davis, 
California 95616.) 


Journal of the Lepidopterists’ Society 
37(2), 1983, 146-147 


THE IDENTITY OF TWO MONOTYPIC GEOMETRID GENERA 
WRONGLY ATTRIBUTED TO THE NEARCTIC FAUNA 
(GEOMETRIDAE) 


DOUGLAS C. FERGUSON 


Systematic Entomology Laboratory, IIBIII, Agricultural Research Service, U.S.D.A., 
% U.S. National Museum of Natural History, Washington, D.C. 20560 


ABSTRACT. Apolema carata (Hulst) and Nyctiphanta laetula Hulst were de- 
scribed from North America in error. A. carata is a synonym of Pseudopanthera en- 
nomosaria (Walker) [Asia], and N. laetula is a synonym of Aspitates ochrearia (Rossi) 
[Europe]. Apolema Hulst and Nyctiphanta Hulst are monotypic genera and junior 
synonyms of Pseudopanthera Hubner and Aspitates Treitschke respectively. 


Apolema carata (Hulst), described from Florida, and Nyctiphanta 
laetula Hulst, described from Arizona, have been consecutively listed 
in all check lists covering North American Geometridae from the time 
the two generic names were proposed (Hulst, 1896:336) to the 
McDunnough check list (1938:168). Apolema carata was listed for 
Florida by Kimball (1965:186); both genera were listed most recently 
by Fletcher (1979:18, 142). I identified A. carata in time to eliminate 
it from the new Check List of the Lepidoptera of America North of 
Mexico (Hodges et al., in press), but not so with N. laetula, which 
was left at the end of the tribe Lithinini, following earlier authors. 
No previous author satisfactorily associated the unique type of either 
species with any known North American geometrid, not surprisingly 
inasmuch as both have proven to be exotic. Through the kind coop- 
eration of Dr. Frederick H. Rindge of the American Museum of Nat- 
ural History, I was able to borrow the types and identify them. 

The holotype of Apolema carata is a specimen of Pseudopanthera 
ennomosaria (Walker, 1862) or something very close to it, probably 
from India or Pakistan. Thus Apolema Hulst, 1896, is a junior syn- 
onym of Pseudopanthera Hubner, 1823, and its type by original des- 
ignation and monotypy, Aspilates carata Hulst, 1887, should be listed 
as a junior synonym of P. ennomosaria (Walker). 

The holotype of Nyctiphanta laetula is a specimen of the European 
Aspitates (Aspilates Auct.) ochrearia (Rossi, 1794). Thus Nyctiphanta 
Hulst, 1896, is a junior synonym of Aspitates Treitschke, 1825, and 
its type by original designation and monotypy, Nyctiphanta laetula 
Hulst, 1896, is a junior synonym of Aspitates ochrearia (Rossi). 

These names should therefore be referred to the synonymy under 
Pseudopanthera and Aspitates as follows: 


VOLUME 37, NUMBER 2 147 


Pseudopanthera Hubner, 1823 


Apolema Hulst, 1896:336. Type-species: Aspilates carata Hulst, 1887: 
211, by original designation and monotypy. Not new synonymy here 
because it was mentioned in my introductory comments in the check 
list (Hodges et al., in press). 


Pseudopanthera ennomosaria (Walker, 1862) 


Aspilates (sic) carata Hulst, 1887:211 (new species based on “1 6, 
Fla. Coll. Franck,” now in the American Museum of Natural His- 
tory). Hulst, 1891:71. 

Apolema carata, Hulst, 1896:336 (new genus proposed for carata). 
Dyar, 1903:315. Smith, 1903:75. Barnes and McDunnough, 1917: 
114. McDunnough, 1938:168. Kimball, 1965:186. Fletcher, 1979: 
18. 


Aspitates Treitschke, 1825 


Nyctiphanta Hulst, 1896:336. Type-species: Nyctiphanta laetula 
Hulst, 1896:336, by original designation and monotypy. NEW SYN- 
ONYMY. 


Aspitates ochrearia (Rossi, 1794) 


Nyctiphanta laetula Hulst, 1896:336 (new genus, new species, de- 
scribed from “Arizona, one male,” now in the American Museum 
of Natural History). Dyar, 1903:315. Smith, 1903:75. Barnes and 
McDunnough, 1917:114. McDunnough, 1938:168. Fletcher, 1979: 
142. NEW SYNONYMY. 


LITERATURE CITED 


BARNES, WM. & J. H. MCDUNNOUGH. 1917. Check List of the Lepidoptera of Boreal 
America. Herald Press, Decatur, Illinois. 392 pp. 

Dyar, H. G. 1902 [1903]. A list of North American Lepidoptera and key to the lit- 
erature of this order of insects. U.S. National Museum Bull. 52:723 pp. 

FLETCHER, D. S. 1979. In I. W. B. Nye. The generic Names of Moths of the World. 
Vol. 3. Geometroidea. British Museum (Nat. Hist.). xx + 243 pp. 

Hust, G. D. 1887. New species of Geometridae, no. 3. Entomol. Americana 2:185-— 
192, 210-212. 

1891. In J. B. Smith. List of the Lepidoptera of Boreal America. American 

Entomol. Soc., Philadelphia. vi + 124 pp. 

1896. Classification of the Geometrina of North America. Trans. American 
Entomol. Soc. 23:245-386, pls. 10, 11. 

KIMBALL, C. P. 1965. Lepidoptera of Florida. Arthropods of Florida and Neighboring 
Land Areas. Vol. 1. Div. of Plant Industry, Florida Dept. Agric., Gainesville. 363 
pp., illus. 

McDunnoucH, J. H. 1938. Check list of the Lepidoptera of Canada and the United 
States of America, part 1, Macrolepidoptera. S. California Acad. Sci. Mem. 1:272 


pp. 
SMITH, J. B. 1903. Check List of the Lepidoptera of Boreal America. American Ento- 
mol. Soc., Philadelphia. vi + 136 pp. 


Journal of the Lepidopterists’ Society 
37(2), 1983, 148-154 


A NEW SPECIES OF SCHINIA (NOCTUIDAE) 
FROM CENTRAL FLORIDA, WITH DESCRIPTION 
OF ITS LIFE HISTORY 


D. F. HARDWICK 
Biosystematics Research Institute, Ottawa, Ontario, Canada K1A 0C6 


ABSTRACT. § Schinia rufipenna, closely related to Schinia tuberculum (Hubner), 
is described as new. The species is a resident of central Florida and feeds in the larval 
stage on Pityopsis graminifolia (Michx.) Nutt. The life history of the new species is 
outlined and the immature stages described. 


While undertaking field work at the Archbold Biological Station 
near Lake Placid, Florida, in the fall of 1979, my wife and I collected 


and reared a new species of Schinia closely related to S. tuberculum 
(Hiibner, 1827). 


Schinia rufipenna, new species 


Description. Eyes full and globular as are those of tuberculum. Antennae filiform 
in both sexes. Inner side of foretibia with two apical spines and two or three additional 
marginal spines; outer side with a single apical spine and one or two marginal spines. 

Maculation similar to that of S. tuberculum (Hbn.) (Figs. 3, 4) but better defined and 
usually with a strong reddish suffusion in both sexes. Species showing a sexual di- 
morphism similar to that of tuberculum with female being smaller, darker and with 
narrower forewings than male. 

Vestiture of head and thorax dark orange, unlike the usual greenish-yellow of tu- 
berculum. Upperside of abdomen black with a yellow band at posterior margin of each 
segment. Underside of body dark yellow or light orange. 

Male with forewing varying from bright reddish-brown to light chocolate-brown, 
without the olive suffusion generally evident on tuberculum, and with crisper mac- 
ulation. 

Transverse anterior line double, pale-filled, broadly excurved, shallowly triarcuate. 
Basal space reddish-brown to light chocolate-brown; usually a pale grey or pale cream 
basal line evident at costal margin. Transverse posterior line usually double, excurved 
around cell, then essentially straight to trailing margin. Median space white to pale 
yellow, variably suffused with reddish-brown to light chocolate-brown; a strongly de- 
fined brown shade along costal margin; inner half of median space usually lightly 
suffused with brown; often the suggestion of a brown median shade. Subterminal space 
concolorous with basal space. Terminal space light orange or white, of variable width. 
Fringe varying from orange to reddish-brown with a series of dark dashes. 

Hind wing black with a narrow yellow outer marginal shade and a yellow median 
band; median band usually divided into two patches by apex of black discal spot. 
Fringe yellow. 

Underside of forewing dark yellow with a basal black patch and a very large subter- 
minal black patch; discal spot variably defined at inner margin of latter; basal and 
submarginal black patches often fused through discal spot, leaving only one or two 
pale median spots or patches. Underside of hind wing dark yellow with a dark patch 
at inner margin, a dark discal spot, and a dark post-median line expanding toward anal 
angle into a broad band. Discal spot often fused proximally with inner patch and distally 
with post-median band. 

Female. Smaller, darker, and with narrower wings than male. Median space more 


VOLUME 37, NUMBER 2 149 


Fics. 14. Schinia spp., Lake Placid, Florida: 1 & 2, S. rufipenna, n. sp., holotype 
and allotype; 3 & 4, S. tuberculum (Hiibner), male and female. 


heavily suffused than in male and usually with white lines evident along the veins. 
Median band of hind wing usually reduced to two rather small yellow spots. 

Mean expanse. Male, 21.3 mm; female, 20.3 mm. 

Male genitalia (Fig. 5). Essentially the same as that of tuberculum, except for the 
conformation of the uncus. Valve elongate and flattened, with a dilated sacculus and 
with a constriction distal to sacculus. Ampulla reduced to a short stub. Corona con- 
sisting of 20 to 25 setae in two or three rows along apical margin of valve. Juxta with 
a high, narrowly rounded dorsal margin and a broadly rounded ventral margin. Uncus 
much stouter apically and mesally than subbasally; uncus of tuberculum slender, only 
slightly stouter distally than subbasally. 

Female genitalia (Fig. 6). Indistinguishable from those of tuberculum; elongate and 
slender. Valve rather short and apically rounded, with a dense clothing of short setae 
and a few elongate ones. Penultimate abdominal segment densely clothed with elon- 
gate slender spicules. 

Type material. HOLOTYPE: ¢ (Fig. 1), Lake Placid, Fla., 8 Nov. 1979, D. and V. 
Hardwick. ALLOTYPE: 2 (Fig. 2), same locality and collectors, 2 Nov. 1979. PARATYPES: 
3 66 and 5 22, same locality and collectors, 1 Nov. to 24 Nov. 1979; 1 2, Orlando, 
Fla., Oct. 1942, D. F. Berry. Holotype and allotype and 8 paratypes in the Canadian 
National Collection (Type No. 16843). One paratype in the collection of the Archbold 
Biological Station. 


Life History and Habits 


Schinia rufipenna occurs sympatrically in central Florida with S. 
tuberculum, and the two species have the same food plant, Pityopsis 
(formerly Heterotheca) graminifolia (Michx.) Nutt. In the immediate 
area in which rufipenna was taken, about six miles west of the town 


150 JOURNAL OF THE:LEPIDOPTERISTS’ SOCIETY 


Fics. 5 & 6. Schinia rufipenna, n. sp., male and female genitalia. 


of Lake Placid, tuberculum was much the commoner species as in- 
dicated by both the number of adults and number of larvae taken. 
The behavior and pattern of development of the two is essentially the 
same. Although the full globular eyes of rufipenna would suggest at 
least partially nocturnal habits, the moths were only active in the 
oviposition containers during the late morning and afternoon, and no 
adults were taken in the light trap during the calendar period that the 
moths were flying. 


VOLUME 37, NUMBER 2 


Fics. 7-10. Ultimate stadium larvae of Schinia spp. on Pityopsis graminifolia (Michx.) 
Nutt.: 7 & 8, S. rufipenna, n. sp., dorsal and lateral; 9 & 10, S. tuberculum (Hubner), 
dorsal and lateral. 


As with most species of heliothentines the adults copulate on the 
flowering heads of the food plant. The female inserts her ovipositor 
into the open head of Pityopsis and deposits the eggs among the 
florets. The newly hatched larva feeds first on the florets and subse- 
quently on the developing seeds. During one of the median stadia 


152 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


the larva moves from one head to a second. The larva continues to 
secrete itself in a Pityopsis head at least until moulting into the last 
stadium. The last instar does not hide in the debris at the base of the 
plant as do many species of the genus but conceals itself along the 
stems of the plant. Its long slender shape probably renders it rela- 
tively inconspicuous in this position. The majority of reared larvae 
matured in five larval stadia, a few in six. The duration of the larval 
stage among reared individuals seemed unusually protracted, its mean 
duration of 37.4 days contrasting strongly with the normal three-week 
period recorded for other species of the genus (see Hardwick, 1958). 

The mature larva burrows into the ground to pupate. The species 
is evidently univoltine, remaining in the pupal stage until the follow- 
ing fall, when its food plant again comes into blossom. 


Immature Stages 


Ege. Pearly white when deposited; remaining unchanged until the day prior to 
hatching when the ocelli and subsequently the whole head capsule gradually become 
visible through the chorion. Egg hatching five to seven days after deposition. 

First instar. Head pale orange-brown, paler than that of tuberculum. Prothoracic 
and suranal shields medium smoky-brown. Trunk white, becoming stained with yellow 
as larva feeds. Mean duration of stadium, 8.1 days. 

Second instar. Head orange suffused dorsally with brown. Ocelli dark brown. Pro- 
thoracic shield smoky-brown with three pale longitudinal shades. Suranal shield smoky- 
grey with a poorly expressed, somewhat darker grey mid-dorsal band and two or three 
evanescent grey subdorsal lines. Mean duration of stadium, 4.5 days. 

Third instar. Head orange-brown with black ocelli; often a pair of blackish arcs 
diverging upward and outward from near apex of frontal triangle. Prothoracic shield 
grey with some brown suffusion mesally; with four black longitudinal bands, the me- 
dian pair converging anteriorly; prothoracic shield usually noticeably elevated above 
cuticular surface of trunk. Suranal shield undistinguished from remainder of trunk. 
Trunk yellowish-grey with brown longitudinal bands. Mid-dorsal band narrow, brown, 
often with a pale median shade. Subdorsal area yellowish-grey with a brown median 
band somewhat paler than mid-dorsal band; in larger specimens, median band with a 
yellow median shade. Supraspiracular area brown with a broad, white or grey, median 
band. Spiracular band pale grey with a white ventral line and often a brown median 
shade. Spiracles dark brown. Suprapodal area pale grey. Mean duration of stadium, 4.5 
days. 

Fourth instar. Head light orange with dark brown ocelli; often a pair of dark brown 
arcs diverging upward and outward from near apex of frontal triangle. Prothoracic 
shield elevated above general surface of trunk; white with four longitudinal black 
bands, the median pair converging anteriorly and occasionally fused along anterior 
margin of shield. Suranal shield undistinguished in maculation from remainder of trunk. 
Mid-dorsal band chocolate-brown with a yellow median shade. Subdorsal area white 
with a median band of paler brown than mid-dorsal band; median band of subdorsal 
area with a dull yellow longitudinal shade. Supraspiracular area brown with a broad 
but discontinuous median white band. Spiracular band white with a weakly expressed, 
multi-arcuate, pale-brown median shade. Spiracles black. Suprapodal area light grey, 
suffused with brown dorsally. Mean duration of stadium, 7.3 days. 

Fifth instar (Figs. 7, 8). Head orange, suffused with brown dorsally; ocelli dark 
brown; a pair of dark-brown arcs diverging upward and outward from near apex of 
frontal triangle. Prothoracic shield prominent, elevated above surface of trunk; white, 


VOLUME 37, NUMBER 2 153 


often suffused with light brown mesally; with four black longitudinal bands, the mesal 
pair converging anteriorly and sometimes fusing along anterior margin of shield. Sur- 
anal shield with maculation undistinguished from remainder of trunk. Overall color of 
trunk orange-brown with a distinctly checkered appearance due to segmental inter- 
ruption of longitudinal banding. Mid-dorsal band pale orange with brown marginal 
lines; marginal lines darker and wider toward anterior margin of each segment. Sub- 
dorsal area white with a dull yellow median band; median band margined with brown 
lines anteriorly on each segment and with orange posteriorly. Supraspiracular area 
consisting of three longitudinal bands: a dorsal brown band fading to orange-brown 
toward posterior margin of each segment; an orange-brown ventral band usually fusing 
with dorsal band at posterior margin of each segment; a discontinuous white median 
band, wide at anterior margin of each segment but narrowing posteriorly and termi- 
nating at fusion of marginal brown bands. Spiracular band white with an inconspicuous, 
multi-arcuate, light-brown median shade. Spiracles black. Suprapodal area pale grey, 
suffused with brown dorsally along margin of spiracular band. Mean duration of sta- 
dium, 12.0 days. 

Pupa. Well sclerotized and somewhat stouter than pupa of tuberculum; orange- 
brown and without characteristic green suffusion of tuberculum. Mesothoracic legs 
relatively long, terminating only a short distance anterior to apex of proboscis. Dorsum 
of fourth abdominal segment with a row of inconspicuous shallow pits. Anterior one- 
third of abdominal segments five to seven slightly elevated above remainder of segment 
and finely pitted; posterior row of pits the most prominent. Rims of spiracles high, 
forming short but definite tubes; spiracles on anterior abdominal segments on a plane 
with general surface of cuticle, those on segments five to seven bore in shallow oval 
pits at margin of raised anterior areas of these segments. Cremaster consisting of four 
spines bome in a single row at apex of a conical projection of tenth abdominal segment, 
the median pair slightly longer and stouter than the lateral pair. 


DISCUSSION 


From the third stadium onward the larva of Schinia rufipenna may 
be distinguished from that of tuberculum by the conformation of the 
two median dark bands of the prothoracic shield; in rufipenna these 
converge toward the anterior margin of the shield, whereas in tuber- 
culum they meet the anterior margin at right angles. In the later stadia 
of rufipenna, the ground color of the trunk is reddish-brown; that of 
tuberculum greyish-mauve. Further, the last instar of rufipenna has 
a somewhat checked appearance because of the segmental interrup- 
tion of the dorso-lateral banding of the trunk; the longitudinal banding 
of tuberculum (Figs. 9, 10) is continuous. 

Schinia tuberculum is widely distributed in the southeastern United 
States. The limits of distribution of S. rufipenna are not known, but 
the species is probably confined to central Florida. The close simi- 
larity in structure and habits of the two species suggests some im- 
mediate common ancestry. During both the Aftonian and Yarmouth 
Interglacials of the Pleistocene, the peninsula of Florida was inun- 
dated by the Okefenokee Sea, and land areas in central Florida were 
reduced to a number of islands in the area now known as the Lake 


Wales Bridge (MacNeil, 1950). 


154 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


The present competition for the same food plant between these two 
closely related species may suggest a fairly recent sympatry. It is 
possible that one population of the immediate ancestor of tuberculum 
and rufipenna evolved into the present day rufipenna during its in- 
sular isolation in the Pleistocene, whereas continental populations 
evolved into the contemporary tuberculum. With the re-establish- 
ment of land connections, tuberculum may have invaded the Florid- 
ian Peninsula resulting in the co-existence of the two species that we 
see today. Certainly, tuberculum is much commoner and more wide- 
spread in the Lake Placid area than is rufipenna. Further, on the basis 
of growth rates in simultaneous larval rearings, tuberculum is the 
more vigorous species. In the face of competition from its more suc- 
cessful relative, rufipenna may well represent a species on the thresh- 
old of extinction. 


ACKNOWLEDGMENTS 


I thank my wife Verna for patient assistance in the field; it was she who took the 
first specimen of rufipenna. I also appreciate the assistance of Mr. Eric Rockburne for 
the preparation of genitalic slides and for assistance with the illustrations accompa- 
nying this paper. 


LITERATURE CITED 


HARDWICK, D. F. 1958. Taxonomy, life history, and habits of the elliptoid-eyed species 
of Schinia (Lepidoptera: Noctuidae) with notes on the Heliothidinae. Can. Ento- 
mol. Suppl. 6:1-116. 

HUBNER, J. 1818-1837. Zutrage zur Sammlung exotischer Schmetterlinge. Augsburg. 

MACNEIL, F.S. 1950. Pleistocene shorelines in Florida and Georgia. U.S. Geol. Surv. 
Prof. Paper 221-F:95-107. 


Journal of the Lepidopterists’ Society 
37(2), 1983, 155-159 


A NEW SPECIES OF EOMICHLA FROM COSTA RICA 
(OECOPHORIDAE) 


J. F. GATES CLARKE 
Department of Entomology, Smithsonian Institution, Washington, D.C. 20560 


ABSTRACT. Eomichla hallwachsae is described as new, is figured and notes on 
its biology are given. 


Eomichla hallwachsae, new species 


(Figs. 1-3) 


Alar expanse. 22-28 mm. 

General description. Labial palpus ochraceous salmon. Antenna ochraceous-salm- 
on; scape light ochraceous-buff. Head vertex light ochraceous-buff; posteriorly light 
buff; face light ochraceous-buff with ochraceous-salmon line laterally. Thorax cream- 
white; base of tegula zinc-orange. Forewing ground color cream-white; from basal 
fourth of dorsum an outwardly curved light clay color, shade extends well into cell and 
reaches termen beyond tornus; in cell and in its outward extremities, this clay color 
shade is heavily overlaid with ochraceous-buff; beyond the clay color shade an out- 
wardly curved line of ground color with a pink tinge; base of costa zinc-orange, this 
color narrowly continued along costa to apex where it broadens to a narrow triangle; 
at apex a short, narrow dash of ground color with a tinge of pink; cilia ochraceous-buff. 
Hindwing ochraceous-buff; veins narrowly outlined with ochraceous-salmon; cilia 
ochraceous-buff. Foreleg mostly zinc-orange, with long light ochraceous-buff and buff 
scales on tibia and tarsal segments; midleg and hindleg mostly ochraceous-buff. Ab- 
domen zinc-orange; abdomen spined. 

Male genitalia (slide USNM 25167). Harpe base of costa with deep excavation; 
sacculus strongly sclerotized; cucullus curved, pointed. Gnathos a sclerotized band 
with lateral, curved extentions. Uncus consisting of two widely divergent arms. Vin- 
culum subtriangular. Tegumen broader than long. Anellus a folded triangular plate. 
Aedeagus stout, with a narrow ridge of short teeth ventrolaterally; vesica unarmed. 

Female genitalia (slide USNM 25168). Ostium transverse, narrow. Antrum broadly 
sclerotized. Inception of ductus seminalis dorsal, slightly before ostium. Ductus bursae 
very short, merging immediately with bursa copulatrix. Bursa copulatrix elongate, 
membranous. Signum absent. 

Holotype. USNM (¢). joc& 4 ¢. 

Type locality. Costa Rica, Guanacaste Prov., Santa Rosa National Park. 

Distribution. Costa Rica. 

Food plant. Bombacopsis quinatum (Jacq.) Dugand. 

Discussion. Described from the ¢ holotype 23-25 May 1980, D. H. Janzen and W. 
Hallwachs, 2 66 paratypes with identical data; 3 63, 2 paratypes same data except 
5-7 June 1980; 5, same except 8-10 June 1980 and ¢ with no date but reared with 
data number 81-SRNP-1089. Paratypes in U.S. National Museum of Natural History 
(USNM) and British Museum (Natural History). 

This species is nearest Eomichla regiella (Busck) but has a whiter thorax and paler 
hindwing. The uncus of hallwachsae has a single arm on each side; the arms of the 
uncus of regiella are divided. The female of regiella is unknown. 

Other species currently placed in the genus Eomichla are: 


E. notandella (Busck), 1911, Proceedings of the United States National Museum, 40: 
209 (Type of Eomichla described in Peleopoda). French Guiana. 

E. nummulata Meyrick, 1916, Exotic Microlepidoptera, 1:546. French Guiana. 

E. thysiarcha Meyrick, 1928, Exotic Microlepidoptera, 3:469. Bolivia. 

E. xystidota Meyrick, 1918, Exotic Microlepidoptera, 2:215. French Guiana. 


156 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


| 


Fic. 1. Eomichla hallwachsae, new species: adult male paratype. 


E. maroniella (Busck), 1911, Proceedings of the United States National Museum, 40: 
208. French Guiana. 


E. imperiella (Busck), 1914, Proceedings of the United States National Museum, 47: 
26. Panama. 


E. irenella (Busck), 1911, Proceedings of the United States National Museum, 40:209. 
French Guiana. 


E. leucoclista Meyrick, 1930, Annalen des Naturhistorischen Museums in Wien, 44: 
231. Brazil. 


Natural History 


The notes on the life history of this taxon which follow were written 
and provided by Dr. D. H. Janzen. 

The only known larval host plant of Eomichla hallwachsae is the 
large native tree Bombacopsis quinatum (Bombacaceae). The large 
palmately compound leaves of B. quinatum have glabrous ovoid leaf- 
lets 7-15 cm long and 5-10 cm wide. The leaf is held in a roughly 
horizontal position and the caterpillar lives on the upper surface of 
the leaflet. The two halves of the leaflet blade are positioned so as to 
form a shallow trough with the leaf midrib running along the bottom; 
the caterpillar spins a dense, white, double-walled tough silk partition 
from one side of the leaflet blade to the other so as to leave a tunnel 
with the dorsal side of the silked-over leaflet midrib as its floor. This 
elongate silk house is 4-8 mm in length initially but becomes as much 


VOLUME 37, NUMBER 2 157 


2 


Fic. 2. Eomichla hallwachsae, new species: ventral view of male genitalia with 
left harpe removed and aedeagus to right. 


as 25 mm long and 6-8 mm wide for the last instar larva. The house 
is positioned roughly in the center of the leaflet. The end toward the 
petiolet is closed, but the other end is always open; the front portion 
of the caterpillar head is visible as a black area well back from the 
tunnel entrance when the caterpillar is not feeding. 

The caterpillar feeds by venturing partly out of the entrance of the 
silk tunnel, both exposed on the leaflet surface and among or under- 
neath a more flimsy layer of silk around the tunnel entrance. It eats 
the surface of the leaflet down to the epidermis of the underside of 
the leaf but leaves the epidermis intact. The result is that, when a 
leaf with Eomichla hallwachsae is viewed from below, there are ir- 
regular ‘windows’ in the leaflet blade around the midrib somewhat 
distal from the center of the leaflet. A caterpillar may spend its entire 
development period on a single leaflet, or it may move to a new leaflet 
if its feeding has removed most of the leaf surface. If it moves it may 
make a new silk tunnel. 

There is never more than one caterpillar per leaflet, but in a case 
of heavy infestation, each of the five leaflets may bear a caterpillar or 
an old silk tunnel. 

When ready to pupate the caterpillar thoroughly reinforces the silk 


158 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Fic. 3. Eomichla hallwachsae, new species: ventral view of female genitalia. 


VOLUME 37, NUMBER 2 159 


of the tunnel and spins a loosely attached tightly fitting dense silk 
trapdoor over the entrance and cuts through the leaf almost all the 
way around the site of the silk tunnel. That portion of the leaf falls 
free, and as the remaining leaf attachments dry, the portion with the 
house-cocoon either falls off the tree into the litter or becomes entan- 
gled with the leaves like any other piece of dead leaf. The caterpillar 
may also spin some silk attachments of the house-cocoon to adjacent 
green leaves, which results in the structure hanging free among the 
leaves. The silk at this stage is rusty brown in color, rather than the 
white that characterized the house when the caterpillar was in it. The 
white silk house looks like old spider webbing; the brown silk of the 
house-cocoon is a close match to dead Bombacopsis quinatum leaves. 
The cocoon-house structure is about 3 cm in length and 6-10 mm in 
cross-sectional diameter. 

The adults emerge about one month after pupation during the rainy 
season, but house-cocoons maintained in dry containers in the labo- 
ratory will produce adults at least two months later when wetted. 
Larvae are common on Bombacopsis quinatum by the second month 
of the rainy season at Santa Rosa (June), and an occasional larva may 
be encountered on the foliage as late as the fifth month of the rainy 
season (September—October). Since B. quinatum is leafless from De- 
cember—January until early May (dry season), the moth probably passes 
the dry season as a dormant pupa. 

Bombacopsis quinatum is a large forest tree with a crown contain- 
ing tens of thousands of leaflets. The larvae of Eomichla hallwachsae 
may be found on the foliage at any point in the crown, but they are 
much more common on outer leaves exposed to the sun and on leaves 
in the upper portion of the crown than on shaded or lower leaves. 
The larvae are never found on young saplings or sucker shoots (1-3 
m tall) and only rarely on well-established small young trees (8-5 m 
in height). In the 1981 rainy season the moths were common; inspec- 
tion of a large B. quinatum crown with binoculars yielded hundreds 
of leaves with the characteristic and conspicuous feeding damage. 
Trees growing in forest and forest edges had conspicuously more lar- 
vae on them than did trees isolated in open pastures. 

The adults come to fluorescent and black lights (mostly males) placed 
at least 800 m from mature B. quinatum. 


ACKNOWLEDGMENTS 


I am indebted to Dr. D. H. Janzen for placing these specimens in my hands, and I 
am pleased to name this species for W. Hallwachs, who was one of the collectors of 
this new species. The drawings were done by Mrs. Elsie Froeschner and the photo- 
graph by Victor E. Kranz, Smithsonian Institution photographer. The study which pro- 
duced the species described above was supported by NSF grant 8-11558. 


Journal of the Lepidopterists’ Society 
37(2), 1983, 160-163 


A BRIEF DESCRIPTION OF THE PHYSIOLOGICAL 
TECHNIQUES—DISC GEL ELECTROPHORESIS, 
INCLUDING GEL PHOTOGRAPHY AND 
THIN LAYER CHROMATOGRAPHY 


THOMAS R. TAYLOR 
P.O. Box 1176, Madison Square Station, New York, New York 10159 


ABSTRACT. Two physiological techniques are described: disc gel electrophoresis, 
including photography of the electrophoresis gels, and thin layer chromatography. 
Modifications of the basic techniques have been worked out which give the best results 
with antennal esterases from the cabbage looper moth, Trichoplusia ni (Hubner). 


The following physiological techniques were developed during 
studies for the M.S. Degree in Entomology at the University of Flor- 
ida, Gainesville. The goal of the research was to study the esterases, 
during late pupal and adult development, in the antennae of the cab- 
bage looper moth, Trichoplusia ni (Hubner) and the role these en- 
zymes may play in pheromone breakdown. Dr. Lee Miller of the Allyn 
Museum of Entomology, Sarasota, Florida, advised me these tech- 
niques may also be of interest to taxonomists. Discussed are the fol- 
lowing methods: disc gel electrophoresis, photography of the gels, 
and thin layer chromatography. 


Disc Gel Electrophoresis 


The basic procedure used was that of Davis (1964). Several addi- 
tions to, or modifications of, this technique were developed: 1, a Sage 
Instrument Syringe Pump, Model 355, was used to layer distilled 
water onto the tops of gels before polymerization so that they would 
have a flat top; 2, the separating gel (1.5 ml) was placed in gel tubes 
of 110 mm length and 4.5 mm internal diameter to give better reso- — 
lution; 3, all solutions were made fresh after 2 weeks, and the am- 
monium persulfate and solution F were made fresh weekly; 4, two 
milliamps of current per gel tube were used initially in the studies, 
but it was later found that 3-4 mamps per tube produced more distinct 
bands; 5, 0.5 ml of 0.001% bromophenol blue, which served as a front 
marker, was added to 50 ml in the upper chamber of Tris-glycine 
buffer, pH 8.3; 6, microsyringes and micropipettes were used to layer 
samples in sucrose through the buffer in the upper chamber onto the 
tops of the stacking gels. 

The electrophoresis run was terminated when the bromophenol 
blue front marker was about 5 mm from the bottom of the gel tube, 
and staining for esterases was carried out immediately. The cabbage 


VOLUME 37, NUMBER 2 161 


a gS a a a a ——— 
/ 
c 4 e822 
c 40 i 
d 
d ed 83 = 2 149 
e:4 Fist cc f 
{ 75 d 133 : 
> d 188 
16 55 
h 
e 175 181 
| 52 x 
33 é a 
n 
° 52 33 
Pp 52 e 
q 64 49 . 
f , 
r 52 j e } 7 55 
f 30 g 99 : 
9 20 
I 
: m 
n 
. cf i ibe 
] 2 3 4 5 6 


Fic. 1. Disc gel electrophoretic comparison of esterases from the antennae of female 
cabbage looper moths with females of five other species of moths (USDA, Gainesville, 
Florida colonies): 1, cabbage looper, Trichoplusia ni (Hubner); 2, fall armyworm, 
Spodoptera frugiperda (J. E. Smith); 3, beet armyworm, Spodoptera exigua (Hubner); 
4, corn earworm, Heliothis zea (Boddie); 5, velvetbean caterpillar, Anticarsia gem- 
matalis (Hubner); 6, Indian meal moth, Plodia interpunctella (Htbner). Letters in- 


dicate bands, while numbers refer to densitometric absorbance, as measured in mm 
from scan. 


looper (Trichoplusia ni) antennal esterases were detected using a- 
naphthyl! acetate and Fast Blue RR Salt, according to the procedure 
of Turunen & Chippendale (1977) with the following modifications: 
1, inhibitors were added to enzyme samples before electrophoresis, 
rather than to the gels following; 2, one ml of acetone was used to 
dissolve the a-naphthyl acetate before addition of phosphate buffer; 
3, the amount of Fast Blue was 3 or 4 mg less than twice the amount 
of a-naphthyl acetate; 4, the phosphate buffer had a pH of 6.8; 5, 
following electrophoresis, the gels were not put into a borate solution, 
but rather were placed directly into a-naphthyl acetate and Fast Blue 
stain; 6, after staining was completed, the gels were stored in the 
following fixative (Mauer, 1971): methanol : water : glacial acetic acid 
(45:45:10). The esterase stain should be made fresh immediately be- 
fore use. 


162 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Gel Photography 


The gels were photographed as soon as they were stained with the 
a-naphthyl acetate and Fast Blue. The gels were left in small glass 
tubes with fixative and were positioned on a light box covered by a 
piece of %” thick opal glass. A circular 22 watt fluorescent light was 
placed 10 cm below the glass. The insides of the box were covered 
with aluminum foil, and the bottom inside was covered with white 
paper. 

A Nikon F-2 camera with a 55 mm Micro Nikor lens was used to 
photograph the gels using Kodak estar base, black and white, SO-115 
(now Technical Pan Film, #2415), shot at !/, second at fll. The film 
was developed at 20°C (68°F) with D-76: water (1:1) for seven min- 
utes. Prints were made on fresh Polycontrast or Kodabrome II, me- 
dium, glossy paper, and were developed in Dektol : water (1:2). Fig. 
1 demonstrates the results of these techniques with a comparison of 
antennal esterases from the female cabbage looper moth, T. ni, and 
five other species of female moths. 


Thin Layer Chromatography 


The cabbage looper pheromone, (Z)-7-dodecene-1-ol acetate, is hy- 
drolyzed into the following products: (Z)-7-dodecene-1l-ol + acetic 
acid. The acetate moiety was tritiated. The two products could be 
separated by the following method. Gelman ITLC-SA, 20 cm x 10 cm 
thin layer paper was cut into pieces 2% cm x 10 cm and ovendried 
at 100°C for 15-20 minutes. Two hundred and fifty ml beakers were 
used as developing chambers, which were covered with glass Petri 
dishes. Seven ml of developing solvent, composed of 15 parts of ethyl 
acetate and 85 parts benzene, were placed in the bottom of the beaker. 
Absorbant paper (Whatman chromatography) was placed against the 
inside wall of the beaker and immersed in the solvent to maintain a 
saturated atmosphere. Ten microliter reaction samples were spotted 
1 cm from the bottom of the plate, and the plates were developed for 
10-15 minutes or until the solvent front was about 1 cm from the top 
of the plate. 

The tritiated acetic acid product stayed at the origin 1 cm from the 
bottom of the plate, while the unreacted pheromone went with the 
solvent front. The alcohol product was unlabeled and not measured 
by this system, although it was verified that the alcohol went up the 
plate to a point between the pheromone and the acetic acid at the 
origin. After development, the plate was cut into sections containing 
the labeled acetic acid and labeled pheromone and counted, using 
Instagel (Packard) in a Packard scintillation counter. 


VOLUME 37, NUMBER 2 163 


ACKNOWLEDGMENTS 


I thank Dr. S. M. Ferkovich, USDA, ARS, Gainesville, Florida, in whose laboratory 
this research was performed, for valuable advice and numerous discussions. 


LITERATURE CITED 


Davis, B. 1964. Disc electrophoresis—II. Method and applications to human serum 
proteins. Ann. N.Y. Acad. Sci. 21:404—-427. 

MAUER, H. R. 1971. Disc electrophoresis and related techniques of polyacrylamide 
gel electrophoresis. Walter de Gruyter, Berlin. Pp. 1-214. 

TURUNEN, S. & G. M. CHIPPENDALE. 1977. Esterase and lipase activity in the midgut 
of Diatraea grandiosella: digestive functions and distribution. Insect Biochem. 7: 


67-71. 


Journal of the Lepidopterists’ Society 
37(2), 1983, 164-165 


GENERAL NOTES 


MYSCELIA ANTHOLIA (NYMPHALIDAE) IN 
THE REPUBLICA DOMINICANA 


According to Riley (1975, Field Guide to the Butterflies of the West Indies, p. 63) 
“there is no real information about the habits or habitat” of Myscelia antholia Godart. 
The species is known only from the Antillean island of Hispaniola and has been re- 
ported from Port-au-Prince, Haiti, and from E] Numero-Azua (1980, Marion Heredia, 
Naturalista Postal, 19/80) in the Republica Dominicana. Thus, there is only one pub- 
lished record of the species from the Republica Dominicana, and indeed there is little 
information available on M. antholia. 

We spent the period between 19 June and 19 August 1981 in the Republica Do- 
minicana, traveling extensively there and making a collection of 1600 butterflies. We 
saw or collected M. antholia at seven localities. Since so little information is available 
on the distribution or habitat of this species, it is appropriate to record our observations, 
which are given in temporal sequence. The first two records are open to some doubt, 
since they are based on individuals which were flying rapidly or which were passed 
while we were traveling in a vehicle. We are confident that these records are based 
on M. antholia, which is unmistakable in the field, but they should be treated with 
some circumspection. The remaining records are indisputable, based on clear sightings 
or specimens collected. 

1) Prov. La Vega, 10 km SE Constanza, Cordillera Central, 1650 m, 2 July 1981. This 
individual was seen by the authors and W. W. Sommer as it crossed an open meadow 
at about 1200 h. The day was bright and sunny and the temperature 30°C. 

2) Prov. La Vega, 3 km SE Constanza, Cordillera Central, ca. 1220 m, 2 July 1981. 
The butterfly was seen from a moving vehicle as it hovered and fluttered above a water- 
filled hole in the road surface at 1300 h. The weather was sunny and bright. The habitat 
was cut-over pine woods and cultivated fields. 

3) Prov. Pedernales, Aceitillar, 35 km NE Cabo Rojo, Sierra de Baoruco, 1220 m, 19 
July 1981. Seen clearly by both authors, the butterfly was flying towards and parallel 
to us along the road of the Alcoa Exploration Company at the bauxite.mine at Aceitillar. 
The area is open pine woods with little shrubby undergrowth. The sighting was at 
1500 h and the temperature was 26°C. 

4) Prov. La Estrelleta, 10 km S Elias Pina, Sierra de Neiba, 732 m, 27 July 1981. 
The butterfly was seen clearly by both authors in a stand of open high-canopied hard- 
woods. When alarmed, the insect flew rapidly up a slope into dense woods and was 
not seen again. The sighting was at 1445 h and the temperature was 36°C. 

5) Prov. San Juan, just SE Sabana Alta, 305 m, 29 July 1981. While we were traveling 
from San Juan to Azua along the Valle de San Juan, a M. antholia was clearly observed 
by both of us as it flew toward and parallel to us in an area of open fields. The habitat 
was generally Acacia scrub, but at the point of observation there was a fencerow cov- 
ered with Antigonon leptopus, by which the butterfly flew without stopping, fled across 
the open field, and was not seen again. The time was 0935 h and the weather was 
bright and sunny. 

6) Prov. Santiago Rodriguez, Loma Leonor, 18 km SW Moncion, northern foothills 
of the Cordillera Central, 550 m, 3 August 1981. One 2 M. antholia was collected in 
moderately dense deciduous riverine woods along the Rio Toma at 1400 h and a tem- 
perature of 38°C. The butterfly was seen flying through the woods with the same erratic 
flight of Hamadryas februa Hubner; when pursued, it fled into the woods but 0.5 h 
later reappeared at the same site as previously and was taken while it rested on a tree 
trunk about 1 m from the base. The insect lit with its wings spread (thus exposing the 
metallic upperside), and then shortly closed them to become inconspicuous, since the 
ventral coloration has a bark-like camouflage. The head was pointed upward while 
resting. The area in the vicinity of the Rio Toma is mixed pine-deciduous woods. 

7) Prov. La Altagracia, 1 km N Playa Bayahibe, sea level, 16 August 1981. Three M. 
antholia were observed and two (1 6, 1 @) collected. The area is semi-xeric lowland 


VOLUME 37, NUMBER 2 165 


forest, quite dense, and all sightings were made along an unpaved road through the 
forest near Playa Bayahibe. The weather had been rainy the previous day (due to the 
passage of Tropical Wave Dennis), and the road was moderately wet and with many 
puddles of standing water. Over one of these, M. antholia was observed by the junior 
author; it had been drinking at 1030 h and was disturbed by the approach of the 
collector. It flew rapidly and erratically, low to the ground (0.5 m) about the collector's 
legs for about 20 seconds, then lit, head down, upon a 2.5 mm diameter sapling at the 
edge of the road very briefly, and when further pursued, flew into the woods. The 
second individual was seen as it flew along the road in a leisurely manner, close to the 
ground. The senior author attempted to catch it, but, alarmed, the butterfly flew some 
10-12 m and came to rest on a tree trunk (0.5 m diameter), about 1.5 m above the 
ground and | m from the road within the woods. The head was pointed down and the 
wings were closed. When approached through the woods, the butterfly flew about 5 m 
further and somewhat deeper into the woods (perhaps 2 m) and landed once again 
with the wings closed. It was collected on this second tree. After the first alarm at the 
attempted capture, the insect flew in a deliberate and non-nervous manner and seemed 
in no hurry to reach the second tree where it rested. The last specimen was observed 
as it flew leisurely along the edge of the road near its opening onto the main paved 
road. It was netted without incident. These observations were made between 1015 and 
1300 h. The weather was very hot and humid. 

To summarize all the above records, in the Republica Dominicana M. antholia occurs 
from sea level to 1650 m but seems more common at higher elevations (above about 
305 m). It is a butterfly of openings in woods (roads and other disturbed areas) into 
which it flies when alarmed. The flight is leisurely when undisturbed, but when pur- 
sued or in widely open areas (sightings 1, 5) it moves with determined and rapid flight. 
Myscelia antholia rests on saplings or trees, often landing with the wings open and 
the head either up or down, remains in this position for about 15 seconds and then 
folds the wings over the back to become inconspicuous. Temperatures of activity range 
between 26°C and 38°C. 

Myscelia antholia may not be so rare as suggested by Riley; it may be rather periodic 
in activity or it may emerge at a time when most collectors are not in the field. The 
number of man-hours (33) spent at locality (7) in 1980 and 1981 without seeing M. 
antholia is truly amazing. Yet most of these hours (19) were in June and July, and these 
months may be too early for M. antholia except for scattered individuals. One other 
fact is pertinent: Of all Dominican localities where we saw or collected M. antholia, 
only (7) has been visited so frequently (6 times). Also, the visit to (7) when three 
individuals were seen was in the late morning; whereas, most other visits have been 
in the early afternoon when the temperatures were higher. It may be that M. antholia, 
like Prepona amphitoe Godart under similar circumstances, becomes inactive and rests 
during the heat of the day (especially in late July and August). 

We wish to thank our fellow collector Kurt M. Iketani for his interest and help, and 
the Alcoa Exploration Company for permitting us to stay at their guest house at Cabo 
Rojo. 


FRANK GALI, 156 S. Melrose Dr., Miami Springs, Florida 33166 AND ALBERT 
SCHWARTZ, Miami-Dade Community College, North Campus, Miami, Florida 33167. 


Journal of the Lepidopterists’ Society 
37(2), 1983, 166-167 


NOTES ON THE AUTUMNAL NORTHWARD MIGRATION OF THE 
CLOUDLESS SULPHUR, PHOEBIS SENNAE (PIERIDAE), 
ALONG THE SOUTH CAROLINA COAST 


On 27 August 1978 large numbers of cloudless sulphurs, Phoebis sennae (Linnaeus), 
were observed moving along the South Carolina coast in a northeasterly direction. The 
butterflies were flying over a salt marsh just inland from Folly Island, a barrier island 
south of Charleston, South Carolina. This flight continued for several days with few 
butterflies stopping at flowers or attempting to oviposit. The flight also was observed 
over grassy areas and fields a few miles inland from the coast, but the flight seemed to 
be most intense just inland from the barrier islands. Ten or more miles inland, the 
cloudless sulphurs seemed to be flying in random directions with no perceptible mi- 
gration taking place. (On 3 October 1978, however, one of us (Laurie) received an 
inland report of cloudless sulphurs moving east in large numbers.) 

On 1 September a procedure was initiated to quantify the continuing migration on 
the immediate coast. A 60 m wide grassy area, bounded on the southeast by a row of 
trees and on the northwest by the offices (located at the South Carolina Marine Re- 
sources Research Institute on James Island, two air miles (3.2 km) south of Charleston, 
S.C.) of the authors, was selected as a study area where daily observations could be 
made. All northerly-migrating cloudless sulphurs passing between the row of trees and 
the windows of our offices flew perpendicular to our line of sight and, therefore, could 
be counted with ease. Five minute counts were conducted at different time periods of 
each day of the migration. The total number of cloudless sulphurs passing through our 
observation area in northerly and southerly directions was recorded for each five min- 
ute period. Counts were made in the morning, at mid-day, and in the afternoon. General 
observations on wind direction, wind speed, and cloud cover were also made. 

During the 1978 migration an early peak was reached on 5 September with an av- 
erage of 125 cloudless sulphurs passing through the count area per five minute period, 
all in a northeasterly direction. The migration continued through 6 October with a 
second peak on 28 September of 323 butterflies moving northeasterly through the 
observation area per five minute count period (64.6 cloudless sulphurs per minute or 
about one per second). The average number of northeasterly-migrating cloudless sul- 
phurs recorded per five minute period in 1978 was 69.5 (n = 29). 

In 1979 a similar migration occurred, beginning again in late August (23 August). 
The same five minute count procedure used in 1978 was employed in 1979 and was 
carried out in the same study area. In 1979 the migration was most intense between 
24 August and 31 August and between 10 September and 9 October. One hundred and 
fifty cloudless sulphurs were recorded during one count period on 31 August, and 160 
butterflies were counted during a five minute period on 9 October. Fewer cloudless 
sulphurs migrated northeasterly during the 1979 migration, the average number of 
butterflies per five minute count period being 48.9 (n = 25). 

Again, in 1980 a northeasterly migration of cloudless sulphurs was observed in the 
study area. The 1980 migration began in late August (28 August) and continued through 
22 September; however, fewer butterflies were seen during the 1980 migration than 
were seen in 1978 or 1979. The average number of cloudless sulphurs observed per 
count period in 1980 was only 15.6 (n = 17) with peak counts of 31 occurring on 11 
and 17 September. 

A review of our general observations on wind speed, wind direction, and cloud cover 
reveals that the migrations were most intense on still, clear days; however, considerable 
flights were recorded on breezy, partly cloudy days. A marked reduction in the number 
of cloudless sulphurs on the wing was obvious on gusty days, with no butterflies flying 
on rainy days. In 1979 Hurricane David passed just south of the study area on 4 Sep- 


VOLUME 37, NUMBER 2 167 


tember, but the northeasterly migration continued on 6 September, after the rains 
associated with the hurricane had passed through the area. 

The northeasterly migration reported herein was limited to late August, September, 
and early October. The cloudless sulphurs flew over marshes and grassy areas and 
generally avoided woodlands. Few cloudless sulphurs were observed flying along bar- 
rier island beaches, as do monarchs (Danaus plexippus (L.)) in their autumnal south- 
ward migration. On the other hand, the cloudless sulphurs migrated in large numbers 
over the salt marshes just inland from the barrier island beaches. In late September 
and early October of 1978 some cloudless sulphurs flew southwesterly, against the 
grain of the larger northeasterly migration. During one count period on 19 September 
1978, 109 cloudless sulphurs were counted flying northeasterly, while 90 were record- 
ed moving southwesterly. The total number of southwesterly-migrating cloudless sul- 
phurs counted during this period could have been influenced by possible northeasterly 
head winds from Charleston harbor—just north of the observation area—that may have 
blown some butterflies back into the study area. During the period of 18 September 
to 6 October 1978 when the southwesterly-migrating cloudless sulphurs were counted, 
an average of slightly less than ten butterflies per five minute count period was re- 
corded (n = 16). 

In his discussion on the cloudless sulphur, Klots (1951, A Field Guide to the But- 
terflies of North America, East of the Great Plains, Houghton Mifflin, Boston, 349 pp.) 
noted that “there appears to be a considerable northward migration in the autumn.” 
In Virginia, Clark and Clark (1951, Smithsonian Misc. Collect. 116(7), 239 pp.) stated 
that they had “kept a special watch” for migratory flights of the cloudless sulphur; 
however, only a few directional flights, all recorded from inland counties, have been 
reported in Virginia. None of these flights were mass migrations; instead, the butterflies 
were widely separated and seldom in sight of each other’ (Clark and Clark, op. cit.). 
Harris (1972, Butterflies of Georgia, University of Oklahoma Press, Norman, 326 pp.) 
pointed out that, in Georgia, the “annual fall migration of the cloudless sulphur ... 
coincides with that of the monarch, ... southeast toward Florida.” In Florida, Walker 
(1978, J. Lepid. Soc. 32:178-190) caught significantly more cloudless sulphurs moving 
southward than northward in Malaise traps set in the autumn of 1975 and of 1976. 

As mentioned earlier, the northward migration of the cloudless sulphur was only 
obvious along the immediate coast, which it seemed to follow northeastward. Inland, 
the butterflies seemed to be moving at random. In light of the fact that there are no 
reports from Virginia, North Carolina, or Georgia to corroborate our South Carolina 
migration data, several basic questions must be asked: Is this migration a local phe- 
nomenon? Or does the autumnal cloudless sulphur migration continue up the Atlantic 
coast? And, why, in autumn, would cloudless sulphurs be flying northward along the 
coast of South Carolina, while in Georgia and Florida the butterflies seem to be moving 
southward? 

Baker (1968, Phil. Trans. Roy. Soc. London, Ser. B, Biol. Sci. 253:309-341; 1969, J. 
Anim. Ecol. 38:703-706) has carefully studied the migrations of British butterflies and 
has developed a theory of the evolution of butterfly migration. He has pointed out that 
most British butterflies have become migratory in response to one or more of the 
following factors: 1) the abundance of predators or parasites; 2) the abundance of a 
preferred food plant; 3) day length; and 4) climate (temperature and/or humidity). The 
combination of the above factors that has resulted in the cloudless sulphur migrating 
northward (in autumn) along the South Carolina coast (and possibly farther north) and 
southward in Georgia and Florida is yet to be explained. 


L. L. Gappy, Rte. 1, Box 223, Walhalla, South Carolina 29691 AND PETE LAURIE, 
612 Flint Street, Charleston, South Carolina 29412. 


Journal of the Lepidopterists’ Society 
37(2), 1983, 168 


ABERRANT HEMILEUCA MAIA (SATURNIIDAE) 


On 1 November 1980 while collecting in Zaleski State Forest in Vinton County, 
Ohio, an albino specimen of Hemileuca maia Drury was taken in flight along the 
margins of a deciduous forest. The location of the capture consists primarily of oaks 
(Quercus spp.), staghorn sumac (Rhus typhina L.), and yellow poplar (Liriodendron 
tulipifera L.). The species is quite common here, and numerous individuals were 
observed and captured along a gravel road which penetrates the forest. The aberrant 
specimen was taken in flight at a height of approximately two meters after emerging 
from the trees just east of the road. The aberrant and typical males were in search of 
concealed females. The specimen (Fig. 1) is very striking in appearence. The normal 
black melanin of the wings is wholly replaced by light beige. The tufts of red scales 
found at the posterior tip of the abdomen and on the thorax are very inconspicuous 
and nearly absent. The white areas of the wings remain but are only slightly apparent 
within the light ground color. The body shares completely the replacement of color, 
providing unquestionable evidence that the specimen is indeed an albino and not 
simply lacking wing scales. 


JOHN V. CALHOUN, 382 Tradewind Ct., Westerville, Ohio 43081. 


Fic. 1. Aberrant ¢ specimen of Hemileuca maia taken in Vinton Co., Ohio, 1 No- 
vember 1980. 


Journal of the Lepidopterists’ Society 
37(2), 1983, 169 


NOCTUA PRONUBA (L.) ON SABLE ISLAND, NOVA SCOTIA, 
A RECORD OF DISPERSAL 


On 22 July 1976 a number of spruce budworm moths, Choristoneura fumiferana 
(Clem.), arrived on Sable Island and were quite active in mid afternoon, flying low 
over the dune vegetation. It was supposed that they must have come from Cape Breton 
Island, 193 km (120 mi.) to the north, where an infestation was in progress. On | 
September 1981 a specimen of Noctua pronuba (L.) was taken in a light trap at the 
Meteorological Station on Sable Island. This time there was no doubt that the moth 
had come from the Halifax area of the mainland, 298 km (185 mi.) to the west. N. 
pronuba, a European cutworm, was first captured in Halifax in 1979 and has since 
spread 66 km (41 mi.) to the southwest and 137 km (85 mi.) to the north (see Map) (B. 
Wright & K. A. Neil, in press). It has not been taken in light traps northeast of the 
Halifax area. 


BARRY WRIGHT, Nova Scotia Museum, 1747 Summer Street, Halifax, Nova Scotia 
B3H 3A6, CANADA. 


de 
Qs 
4 


Cape Breton 


Island 


Map. Noctua pronuba distribution in Nova Scotia, 1981. 


Journal of the Lepidopterists’ nee 
37(2), 1983, 170-171 


THE SECOND SPECIMEN OF EPARGYREUS SPANNA (HESPERIIDAE) 


Epargyreus spanna was named (Evans, 1952, Cat. Amer. Hesperiidae, Pt. I1:46) on 
the basis of a single female with the locality datum of “Santo Domingo’; the specimen 
had been collected in 1855. Riley (1975, Field Guide to the Butterflies of the West 
Indies, p. 158) characterized and figured (PI. 21) the species. Epargyreus spanna has 
apparently not been collected since 1855. Brown and Heineman (1972, Jamaica and 
Its Butterflies, p. 357) suggested that E. spanna is-a subspecies of Jamaican E. antaeus 
Hewitson; although the two species are similar in details of the female genitalia and 
in having a bold silvery-white bar on the unhw, E. antaeus is much the smaller of the 
two taxa (female fw length 27-28 mm in antaeus, 34 mm in spanna). Both species 
appear to be rare (Brown and Heineman listed only thirteen localities for E. antaeus, 
and Evans examined only 21 specimens), but E. spanna has been known only from 
the holotype. 

The specimen is a gravid female with a fw length of 35 mm. The up ground color of 
both wings is very dark brown, the uphw unmarked, the upfw with three subapical 
white dots, the lowest one in R;—M, displaced more apically than those in R,—-R,; and 
R,-R, and comma-shaped. The remaining five spots are yellowish and arranged as in 
Riley’s color plate except that the lowermost in Cu,—2A slightly overlaps that in Cu,— 
Cu,, and that in the cell is not indented along its outer margin. The unfw is marked 
and colored like the up. The silvery-white hw bar is 5 mm wide at its widest part and 
the short brown bar that breaks the costal extreme of the white bar almost completely 
cuts off the more discal portion of the white bar at this costal margin. Riley also showed 
a pale semilunar marking at the anal end of the white bar; this pale marking is barely 
discernible in our specimen (Fig. 1). 

The area where the E. spanna was taken lies at 915 m and 10 km W Jayaco, Provincia 
de la Vega. The site is a mountain torrent, strewn with boulders and alternating with 
flat areas of slack water. The stream is reached by a 100 m path from the Jayaco—E]l 
Rio road. The region is generally well forested with deciduous forest, in which the 
forest palm (Prestoea montana) is common. The stream is generally open (i.e., the 
canopy does not close above it), and the forest may come to the stream edge, or the 
banks may have a border of shrubs, bushes, and grasses. 

The skipper was taken at 1400 h on 17 August 1981. The precise area was along a 
slack-water pool about 0.3 m deep; below the pool the stream was steep and torrential, 


Fic. 1. Upper and underside views of female E. spanna (7327 in the collection of 
the junior author). 


VOLUME 37, NUMBER 2 Zl 


above the pool a steep (and impassable) fall. The stream at this point is about 5 m wide 
and is open, with a high canopy on the sloping banks, but with shrubs and bushes 
along the stream itself. The day was overcast in general, but the sun alternately ap- 
peared and disappeared. Generally, collecting was poor and at the site of capture of 
the E. spanna no other butterflies were seen. Just below the pool Lycorea ceres Cramer 
was moderately abundant but not (the usually common) Greta diaphana Drury. The 
estimated distance of this spot is 1 km from the path from the road to the stream, but 
this estimate may be too great. In any event, the pool is at the upper end of the stream 
beyond which it is difficult to continue. 

The E. spanna flew across the stream; the flight was slow and ponderous and not 
darting, perhaps due to the greatly enlarged abdomen with eggs. The skipper landed 
on the top of a leaf of a streamside shrub about 1.5 m above the ground and adjacent 
to the pool; the wings were held open. The general impression of the skipper in flight 
was of Colobura dirce Linnaeus (doubtless due to the white underside pattern), but 
the flight was completely different from the rapid and darting flight of that species. 

We are grateful to Kurt M. Iketani for his companionship and for taking the photo-— 
graphs of the specimen. 


FRANK GALI, 156 S. Melrose Dr., Miami Springs, Florida 33166 AND ALBERT 
SCHWARTZ, Miami-Dade Community College, North Campus, Miami, Florida 33167. 


Journal of the Lepidopterists’ Society 
37(2), 1983, 171-174 


BATTUS ZETIDES IN THE REPUBLICA DOMINICANA 


The papilionid Battus zetides Munroe is endemic to the Antillean island of Hispan- 
iola. Hall (1925, The Entomol. 58:162) considered the species “Apparently very rare” 
and recorded a single specimen from La Vega, Republica Dominicana as the only 
example with “exact” locality data, although there existed other specimens (with im- 
precise locality data—Haiti—in British collections). Riley (1975, Field Guide to the 
Butterflies of the West Indies, p. 140) gave the range in an anomalous manner: “Known 
only from Haiti, La Vega, and apparently very rare. Should also occur in the Dominican 
Republic.” His statement of range is, of course, taken from Hall, but he has confused 
the two countries involved. Marion Heredia (1980, Naturalista Postal, 26/80) noted the 
capture of a series of specimens on 3 October 1976 at Las Auyamas, Polo, Provincia 
de Barahona, Republica Dominicana; he regarded these as the first specimens from 
that country, apparently unaware that Hall had mentioned the La Vega locality many 
years earlier. Considering the time of that record, it seems likely that it does not apply 
to the city of that name which lies at an elevation of about 100 m, but rather to the 
Cordillera Central south of La Vega. The Las Auyamas record is from an elevation of 
about 1000 m. 

Additionally, Riley (1975:P1. 18) figured a specimen of B. zetides without tails; whereas, 
Lewis (1973, Butterflies of the World, p. 23) figured a specimen with tails. This can be 
clarified immediately, since the plate in Riley is in error; the species is indeed tailed. 

Between 19 June and 19 August 1981, we made collections of butterflies throughout 
much of the Republica Dominicana. The period of 13 July to 21 July we spent at the 
guest house of the Alcoa Exploration Company at Cabo Rojo. In this region there are 
two roads that ascend the mountains to the north: 1) Alcoa’s private road to their bauxite 
mines at Aceitillar in the Sierra de Baoruco, and 2) the Dominican border road that 
parallels the Dominico-Haitian border from Pedernales (at sea level) to Los Arroyos 
on the southern face, and thence over the mountains (which here are a continuation of 


2 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


the Haitian Massif de la Selle) to El Aguacate on the northern face. The maximum 
elevation reached by the Alcoa road is about 1220 m, that of the border road about 
2362 m. Both offer an excellent vertical transect of the ecology of the region, going 
from xeric desert to mesic deciduous forest to upland pine (Pinus occidentalis) forest. 

We were able, with the use of a jeep supplied us through the offices of Dr. Eugenio 
de J. Marcano of the Museo Nacional de Historia Natural de Santo Domingo, to reach 
Los Arroyos on the border road. This village and army post are at an elevation of about 
1100 m. Much of the southern slope of the Massif de la Selle through which the border 
road passes was formerly rich mesic deciduous forest, but now most of this forest has 
been cut, and sloping fields and a few cafetales are the dominant vegetational features 
today. The road becomes barely passable above Los Arroyos, and on the day (15 July) 
that we reached that settlement, the higher slopes were already covered with heavy 
mist and drizzle at 1130 h, and it seemed pointless to proceed further. The junior author 
has crossed the main ridge between Los Arroyos and El] Aguacate and formerly (and 
presumably even now) one reaches at high elevations (2300 m) a pine-clad plateau 
after passing through a wide horizontal band of deciduous forest within which Los 
Arroyos lies. 

The Alcoa road, when it ascends the southern front range of the Sierra de Baoruco, 
reaches semi-mesic deciduous woods (as at Las Mercedes) which at about 915 m give 
way to pines. This pine forest continues upward to elevations of at least 1220 m; it is 
open, with grassy and shrubby undergrowth and in general appearance is unproductive. 
The road stops at the current bauxite mine at Aceitillar, but a dirt-and-rock road pro- 
ceeds from there at least 12 km to the northwest and an elevation of 1130 m. The road 
passes several abandoned experimental mines and leads to a site called Las Abejas. 
Las Abejas is unique in this region, since it is upland mesic deciduous forest, appar- 
ently totally surrounded by pine forest: thus, a local enclave of deciduous forest in 
what is otherwise pine forest. One comes upon the Las Abejas area abruptly; the road 
has been traveling through pines at elevations of between 1100 and 1220 m, and then 
descends within a kilometer to 1130 m, in beautiful hardwood forest. The slopes are 
steep, and most collecting was done along the margin of the road; at Las Abejas itself, 
there is a wide and fairly level path that leads through the forest. Butterflies were 
abundant both along the road and within the forest, although the species diversity is 
rather limited. The region is seldom visited, and the forest seems relatively uncut. 
Because of the elevation, most profitable collecting must be done in the morning or 
early afternoon, since later in the day mist and drizzle or rain with overcast skies 
regularly brings an abrupt end to collecting. 

On our first visit to Las Abejas on 18 July, we collected between 1130 and 1515 h; 
the temperature was 30°C, the weather alternately sunny and overcast. We collected 
along the sloping road noted above but not on the path through the forest. The senior 
author, within minutes of the beginning of our collecting, netted a B. zetides as it flew 
across the road. Several others were seen at this precise point, leisurely circling the 
tops of vine-covered trees about 10 m high and thus inaccessible. But we soon discov- 
ered that walking along the road up the slope was a ready source of specimens, where 
the butterflies were flying leisurely (but not seen feeding on the moderately abundant 
flowers). The insects seemed undisturbed by the presence of the collectors and were 
not unduly frightened (as are some papilionids) when attempts were made to catch 
them (i.e., there was no alarm behavior). 

On a second visit on 19 July, we collected between 0900 and 1415 h; the temperature 
was 28°C and the weather was generally overcast and sunny in the morning, and over- 
cast in the afternoon. On this occasion we collected not only along the road but also 
on the path through the forest. Along the latter, a few B. zetides were seen, especially 
in areas where there were openings or clearings, rarely within the woods proper. Along 
the road itself, we estimated that we saw 50 individuals. In fact, B. zetides is certainly 
at this locality a very common species of butterfly, if not the most common. 

Our third visit was on 20 July, between 1430 and 1530 h. The day was cool and had 
been heavily overcast; only two B. zetides were seen and none collected. In fact, 


VOLUME 37, NUMBER 2 . eS 


Fic. 1. Upper and lower views of male Battus zetides (6326 in collection of junior 


author). 


butterfly activity in general was reduced to a bare minimum, both on the road and the 
path. 

A total of 10 B. zetides was secured. Most of these were taken along the road where 
they were abundant. We have the impression that B. zetides is a butterfly of dense 
woods, but that it flies in sunny open areas (as along roads or in clearings in the forest). 
The flight is slow and deliberate and generally rather high (3.5 m) above the ground, 
but the butterflies descend to lower levels (2 m) or may fly as high as 10 m with some 
degree of regularity. Securing specimens is not difficult, but the collector must be 
patient and await descent of the butterflies to levels within reach. Attempting to enter 
the forest on steep slopes to catch an individual rarely met with success. 

The series consists of six males (one of which is slightly flown) and four females 
(three of which are slightly to well flown). Contrary to Riley’s (Pl. 18) illustration, the 
upperside band is not unicolorous on both wings; the upperside forewing band (upfw) 
is paler (Pl. 11L8; color designations from Maerz and Paul, 1950, A Dictionary of Color) 
than that of the upperside hind wing (uphw) (PI. 11111) which may be more orange 
(Pl. 10L12) in fresh specimens of both sexes (see Fig. 1). More flown specimens of 
both sexes have the up bands much more yellow, but even in these individuals the fw 
band is paler (Pl. 10G2) than the hw band (PI. 10L4). In fresh specimens, the up both 
wings is a rich blackish chocolate. The us markings of both wings are as figured by 
Riley, and the silvery arrowhead-shaped markings stand in bold contrast to the yellow, 
red, and black ushw markings and ground color. Forewing lengths in males vary be- 
tween 35 and 40 mm (x = 36.0) and in females between 39 and 44 mm (x = 42.0); thus, 
there is some sexual dimorphism in size, but it is not striking. 

It is pertinent to comment on the apparent local occurrence of this species. We also 
collected less than a kilometer southeast of Las Abejas. The area was pine woods, but 
we found a narrow run-off ravine which had a local stand of deciduous trees and a 
dense understory of blackberries (Rubus sp.). The elevation was 1220 m, thus slightly 
higher than Las Abejas, and at the top of the hillslope along which we collected and 
observed such an abundance of B. zetides. We collected at this second locality on 14 
July (0930-1330 h; T = 30°C; bright and sunny in morning, overcast after 1200 h) and 
on 18 July (0920-1115 h; T = 23°C; weather alternately sunny and overcast). No B. 
zetides were seen at this locality, although there was a great deal of local butterfly 
activity both in the pines and in the ravine. Note that our second visit to this locality 
(16 July) was the same day as our first visit to Las Abejas, where B. zetides was common. 
Although there may be a variety of factors at work, certainly the most obvious is the 
richness and much greater areal extent of the forest at Las Abejas in contrast to the 


174 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


narrow stand of deciduous trees at the ravine. It may well be that B. zetides demands 
extensive stands of forest. 

It may also be instructive to compare the old La Vega locality and Mari6n’s Las 
Auyamas locality with Las Abejas. The former lies much lower (assuming that the 
specimen(s) came from La Vega itself, an unlikely possibility) but is in a basically 
mesic area (presently much cultivated but with cafetales and cacaotales and their high- 
canopy shade-trees). If the La Vega material came from south of that city on the north- 
ern slopes of the Cordillera Central, these slopes today are open pine woods with some 
deciduous forest in wide ravines (as below Buena Vista). The latter may well be or 
have been satisfactory for B. zetides, but we visited this area in June 1981 and saw no 
individuals. Las Auyamas on the other hand is in the uplands of the Sierra de Baoruco. 
This area presumably was once well forested, since there are still extensive cafetales 
with their shade trees present (=pseudoforest). The junior author visited the Las Au- 
yamas area on 4-6 August 1980, but the weather conditions, due to the passage of 
Hurricane Allen, were not propitious for butterfly collecting. Still, the area about Las 
Auyamas seems suitable for B. zetides and rather comparable to that at Las Abejas. 

We are grateful to the staff of the Alcoa Exploration Company, especially to Sr. 
Alfredo Lebron and Sr. Victor Garcia, for allowing us to stay at their facility at Cabo 
Rojo; without the loan of the jeep from the Museo Nacional under the directorship of 
Dr. Marcano, the trip to Las Abejas would have been more arduous, and we are grateful 
to him and his staff for their assistance. The illustration is the work of Kurt M. Iketani; 
we acknowledge his efforts with pleasure. 


FRANK GALI, 156 S. Melrose Dr., Miami Springs, Florida 33166 AND ALBERT 
SCHWARTZ, Miami-Dade Community College, North Campus, Miami, Florida 33167. 


Journal of the Lepidopterists’ Society 
37(2), 1983, 174-176 


PUPAL SIZE AND EGG PRODUCTION CHARACTERISTICS IN 
ROTHSCHILDIA FORBESI (SATURNIIDAE) 


Rothschildia forbesi Benjamin occurs in the United States in the Rio Grande Valley, 
Texas. According to Ferguson (in R. B. Dominick et al., 1972, The Moths of America 
North of Mexico, fasc. 20.2B, Bombycoidea) there is only limited information available 
on the biology and early stages of this species. During 1981 I used a series of 23 wild 
R. forbesi pupae, and five of the subsequent adult females to collect data for methods 
development modeling. The R. forbesi were used simply because they were available 
for study at the time a modeling data set was needed. However, the data that were 
collected, aside from being used for methods development research, provide funda- 
mental information on pupal dimensions and egg production characteristics for this 
little-studied species. 

Pupal sex was determined by examining the genital openings, which were very 
distinctive in the R. forbesi pupae studied. Males had a single opening on the venter 
of the 9th abdominal segment; and females had single openings on the venter of the 
8th and the 9th abdominal segments. All pupal sex determinations were confirmed at 
adult emergence. Pupal weights and dimensions, determined as described by Miller 
et al. (1982, J. Lepid. Soc. 36:207—216), are summarized in Table 1. Mosher (1916, Ann. 
Entomol. Soc. Amer. 9:136—158) described pupae of Rothschildia orizaba (Westwood) 
and R. cincta cincta (Tepper) (the latter under the name R. jorulla; see C. Lemaire, 
1978, Les Attacidae Americains, Attacinae, Neuilly-sur-Seine, France), indicating that 
R. orizaba pupae were 23-27 mm in length and about 50 mm in circumference; while 


VOLUME 37, NUMBER 2 175 


TABLE 1. Weights and dimensions of Rothschildia forbesi pupae. 


Mean =: S.D. 

Measurement! Male (n = 12) Female (n = 11) 
Weight BUG 28 OLD 3.20 + 0.48 
Length 27.65 = 25 29.85 + 1.73 
Width 11.20 + 0.39 13.70 + 0.81 
Circumference Shae OUST 45.45 + 2.98 
Antenna length LESS) == Oro 13.08 + 0.99 
Antenna width 3.50 + 0.52 3.09 + 0.30 
Antenna length to width ratio Selo) a= LOS 4.23 + 0.46 


1 Weights in g; dimensions in mm. 


R. jorulla pupae were 25-28 mm in length and about 45 mm in circumference. She 
did not discuss R. forbesi pupae, nor did she state the numbers or sexes of pupae of 
the other Rothschildia species she examined. The information presented here for R. 
forbesi (Table 1), however, is in general agreement with the sizes reported by Mosher 
(1916) for the other species of Rothschildia. 


100 
ra) 
Ww 
= 
” 
Oo 80 
o 
Ww 
a 
7) 
SO 
os) 60 
Ww 
ve 
fo) 
fe 
Ww 
a 40 
= 
= 
z 
z 
< 
Ww 
Ss 20 
(0) 


1 2 3 4 5 6 7 8 9 
NIGHTS AFTER MATING 


Fic. 1. Oviposition pattern for Rothschildia forbesi females. 


176 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Five of the females that emerged were mated, and eggs were collected in paper bags 
each night thereafter until death occurred. Later, the bags were cut open and all eggs 
were counted. To determine total egg complement the abdomen of each female was 
dissected after death and eggs remaining in the ovaries were counted. The average egg 
complement was 232.40 + 42.76; the average number of eggs deposited was 216.2 + 
53.14. Thus, on a percentage basis the females deposited an average of 92.95 + 13.48 
percent of the eggs they emerged with. The only known information on R. forbesi egg 
production is the report by Collins and Weast (1961, Wild Silk Moths of the United 
States, Collins Radio Co., Cedar Rapids) that one female laid “all” of her eggs (156) in 
one night. In the R. forbesi studied here, the average longevity after mating was 7.80 + 
0.84 nights; most of the deposited eggs (>80 percent) were laid during the first three 
nights after mating. None of the females deposited all eggs in any one night. The three- 
night average for eggs was 57.92 + 11.42. There was a positive correlation between 
egg complement and the three-night average for eggs (r = 0.70), which is described by 
the following regression equation, where Ey, = three-night average eggs and FE, = egg 
complement: 


Figs = 25.35 ate 0.15F; 


The oviposition pattern for R. forbesi (Fig. 1) is similar to patterns known for other 
giant silkworm moths (Hyalophora cecropia (L.), Tashenberg & Roelofs, 1970, Ann. 
Entomol. Soc. Amer. 63:107—-111; Hyalophora gloveri gloveri (Strecker), Miller, 1978, 
J. Lepid. Soc. 32:233-234; Callosamia promethea (Drury), Miller & Cooper, 1977, J. 
Lepid. Soc. 31:282-283; Antheraea polyphemus (Cramer), Miller & Cooper, 1980, J. 
Lepid. Soc. 34:256—259). 

The author acknowledges with gratitude the assistance of R. S. Peigler, who collected 
the Rothschildia forbesi pupae at Harlingen, Texas, in January 1981. The opinions 
contained herein are those of the author and should not be construed as official or 
reflecting the views of the Department of the Army. 


THOMAS A. MILLER, U.S. Army Medical Bioengineering R&D Laboratory, Fort De- 
trick, Maryland 21701. 


Journal of the Lepidopterists’ Society 
37(2), 1983, 176-177 


OCCURRENCE OF MEGISTO CYMELA (SATYRIDAE) AT FLOWERS, 
WITH A BEHAVIORAL NOTE 


Most Satyridae are thought not to normally utilize nectar sources (Emmel, 1975, in 
Howe (ed.), The Butterflies of North America, p. 80). Megisto cymela (Cramer) to my 
knowledge has never been recorded visiting flowers. On 9 July 1980 and 5 July 1981 
I observed repeated nectaring by this species on staghorn sumac, Rhus typhina L., in 
Philadelphia, Pennsylvania. 

The habitat is a burn area dominated by the grass Andropogon scoparius. There are 
many clumps of trees and shrubs invading this area, such as gray birch (Betula popu- 
lifolia), bigtooth aspen (Populus grandidentata), hawthorns (Crataegus spp.), cherries 
(Prunus spp.), and staghorn sumac (Rhus typhina). It is surrounded by a climax Tran- 
sition Zone woodland which is part of the Wissahickon Creek Ravine in Fairmount 
Park, Philadelphia. This burn scar, where the butterflies were seen, is actually at the 
top of part of this ravine about 104 m above sea level. 

Megisto cymela is univoltine here, emerging in mid or late June, with worn individ- 
uals being found in August. These common butterflies are usually found flying near 
the ground in their characteristic weak dancing or skipping manner, moving in and out 


VOLUME 37, NUMBER 2 IEEE 


of shrubs or thickets of small trees. On 9 July 1980 a single individual was seen nec- 
taring on the yellow-green inflorescence of Rhus typhina. On 5 July 1981 at 1400 h, 
two individuals were seen nectaring on this flower species in a shaded thicket. The 
day was cloudy and very humid with the air temperature about 29°C. The first indi- 
vidual was observed for over 15 min, moving slowly from one blossom to another before 
disappearing out of view. The second one was found in another clump of these trees 
but nectared at the flowers only briefly. The first butterfly seen on 5 July 1981 had 
initially been found resting on the leaves of Rhus typhina with its wings open and flat, 
very much like a geometrid moth. Later, another individual was also found resting on 
leaves of these trees in this manner, certainly very uncharacteristic of members of this 
family. 

Other butterflies of interest found in this area include Parrhasius m-album (Bdv. & 
LeConte), Satyrium liparops (LeConte), Harkenclenus titus (F.), Atrytonopsis hianna 
(Scudder), and Hesperia metea Scudder. The latter, however, may have been recently 
extirpated here. 


THOMAS S. WILLIAMS, 2366 Rosemore Ave., Glenside, Pennsylvania 19038. 


Journal of the Lepidopterists’ Society 
37(2), 1983, 177-179 


THE “WHITE MALE” VARIANT OF COLIAS (PIERIDAE): 
TWO NEW RECORDS FROM COLORADO 


Male Colias butterflies with their ground color white or near-white, in contrast to 
the typical yellow and orange phenotypes, are extremely rare. Such “white males” 
have been reported in at least seven Colias species (see review by Remington, 1954, 
Adv. Genetics 6:403-450). Wild captured white males are few; they occasionally seg- 
regate out of inbred laboratory strains and mass cultures of these pierids. Here I report 
captures of two more white male Colias, one being from a species in which this variant 
has not previously been recorded. 

On 8 July 1977 I collected a white male C. meadii Edw. (Fig. 1) at the Mesa Seco, 
elev. 3590 m, 8 km west of Lake City, Hinsdale County, Colorado. I am not aware of 
other captures of white males for this Colias species. The specimen was initially mis- 
taken for an “alba” female as it flew down a steep grade. C. meadii “alba” females are 
themselves uncommon in Colorado (Remington, 1958, Proc. X Intl. Congr. Entomol. 
2:787-805; Ferris, 1972, Bull. Allyn Museum 5:1—23) and have never been captured at 
Mesa Seco during a decade of mark-recapture studies by Watt’s Stanford research group 
(W. B. Watt, pers. comm.). 

I also collected a white male C. philodice eriphyle Edw. (Fig. 2) on 3 August 1977 
in an alfalfa field near State Route 92, elev. 1645 m, 8 km west of Hotchkiss, Delta 
County, Colorado. The “alba” phenotype frequency in C. p. eriphyle females in some 
of these agricultural populations is in the neighborhood of 15 percent or less. Here, as 
in much of North America, positive identification of some white females to species is 
hampered by the presence of migrant C. eurytheme Bdv. (whose “alba” frequencies 
in western Colorado are generally below 10 percent) and concomitant introgression. 
Rearings of “alba” females from pure yellow C. p. eriphyle (and the reciprocal) taken 
in fields near Montrose, Colorado, demonstrate that “alba” does occur in pure C. p. 
eriphyle and not just as a result of introgression with C. eurytheme. 

It should be noted that the coloration of “white male” Colias differs significantly 
from that of their white female counterparts. White males, and some of the white 


178 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Fics. 1, 2. White male Colias: 1, specimens of Colias meadii; top: normal male; 
bottom: white male; 2, specimens of Colias philodice eriphyle; top: normal male; 
bottom: white male. Locality data given in text. Color filter used to enhance contrast 
between white males and normal males. 


females, are products of autosomal alleles recessive to others for typical ground col- 
oration (e.g., the genes “whitish” and “blonde,’ Remington, 1954, Lepid. News 7:139- 
145). These characters are not sex-limited, as is “alba,” but itis believed that “whitish” 
and “blonde” females generally pass unnoticed due to their phenotypic similarity to 


Fic. 3. Specimens of Colias alexandra: Left, male “black-vein” variant; Right, 
normal male. Locality data given in text. Compare “black-vein” C. alexandra to the 
white male and “black-vein” C. meadii in Fig. 1. 


VOLUME 37, NUMBER 2 179 


the vastly more abundant “alba” females. For recent reviews of the biochemistry and 
adaptive value of the “alba” variant see Watt (1973, Evolution 27:537-548) and Graham 
et al. (1980, Proc. Natl. Acad. Sci. USA 77:3615—3619). The selective (if any) and bio- 
chemical details of the white male coloration remain unknown. 

The new white male C. meadii also exhibits a second interesting genetic character, 
that of “black-vein” (Fig. 1 does not show this character particularly well). A typical 
and a wild-captured “black-vein” C. alexandra Edw., both taken 5 km east of Crested 
Butte, elev. 8950 m, late June 1977, are shown for comparison (Fig. 3). Ae (1958, 
Genetics 43:564-576) demonstrated that “black-vein” is almost certainly the product 
of a single autosomal allele. The white male C. meadii is indeed curious, as the viability 
and/or penetrance of the “black-vein” character appear low (ibid.; Remington, op. cit.). 
The two white males have been deposited in the entomological collections at the 
Peabody Museum of Natural History, Yale University. 


LAWRENCE F. GALL, Department of Biology, Yale University, New Haven, Con- 
necticut 06520. 


Journal of the Lepidopterists’ Society 
37(2), 1983, 179-180 


ON THE STATUS OF PSEUDOTHYATIRA EXPULTRIX (GRT.) AND 
EUTHYATIRA PENNSYLVANICA J. B. SMITH (THYATIRIDAE) 


Pseudothyatira cymatophoroides (Guenée, 1852) and P. expultrix (Grote, 1863) were 
described as distinct species and continued to be regarded as such until about 1917. I 
am not sure who was responsible for the change, but in the Barnes & McDunnough 
check list of that year expultrix was treated as a form of cymatophoroides. There it 
remained until 1966, when Werny, in a world revision of a large part of the Thyatiridae, 
restored it to specific rank (p. 322), citing in support of this some minor genital differ- 
ences as well as the more obvious differences in wing markings. I have dissections of 
several specimens of each form and can see no differences in the genitalia. The two 
“species” always occur together, from Newfoundland to British Columbia, south to 
northern California, Maryland, West Virginia, Kansas, and in the Appalachians to North 
Carolina (probably, also the White Mountains, Arizona, but only one example seen, a 
male of the nominate form from Pinetop, Navajo Co., about 8000 ft, R. B. Nagle col- 
lection). I have recently seen both forms from a locality much farther south than pre- 
viously reported—West Feliciana Parish, Louisiana (V. A. Brou collection). It is, there- 
fore, not surprising that doubts concerning the validity of Werny’s taxonomic change 
should have persisted. I know that these moths have been reared by others, but no 
conclusive results of such a test have appeared in the literature. 

On 31 May 1980 I collected at bait a female of the nominate (well-marked) form (Fig. 
1) at Colesville, Montgomery Co., Maryland, and from eggs laid by this moth reared a 
brood of 37 adult progeny in August and September of the same year. The larvae were 
reared on Betula nigra L., B. populifolia Marsh, and Prunus virginiana L., as available. 
Sixteen of the offspring were of the nominate form (Fig. 2), and 21 were of form ex- 
pultrix (Fig. 3), showing conclusively that these are indeed forms of the same species. 

The situation with respect to Euthyatira pudens (Guenée, 1852) and E. pennsylvan- 
ica J. B. Smith, 1902 is not so certain. Werny (1966, Untersuchungen tber die Syste- 
matik der Tribus Thyatirini, Macrothyatirini, Habrosynini und Tetheini (Lepidoptera: 
Thyatiridae), Inaugural-Dissertation, Universitat des Saarlandes, Saarbriicken, Ger- 
many, pp. 237, 245) also elevated pennsylvanica from the status of an infrasubspecific 
form to that of a species. The few pudens that have been reared from eggs have turned 


180 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Fics. 1-3. Pseudothyatira cymatophoroides: 1, 2, Colesville, Montgomery Co., 
Maryland, 31 May 1980, parent of brood; 2, 6, reared from ? shown in Fig. 1; 3, 3 
of form “expultrix,” reared from 2 shown in Fig. 1. About natural size. 


out to be of the same form as the parent, which does not prove anything. There is still 
a need for broods to be reared from females of pennsylvanica. The circumstances are 
different from those of E. cymatophoroides in that normal pudens has a very wide 
distribution similar to that of cymatophoroides; whereas, pennsylvanica seems limited 
to certain areas of the Middle Atlantic States. This is the region where industrial mel- 
anism has been most prevalent in North America, affecting perhaps as many as a hundred 
species, and I had supposed that pennsylvanica was the industrial melanic of pudens. 

Werny (p. 245) also introduced a problem of authorship with respect to the name 
pennsylvanica. In elevating the name to specific rank, he listed himself as author in 
accordance with Article 10b of the International Code of Zoological Nomenclature. 
Although Smith (1902, J. N.Y. Entomol. Soc. 10:34) referred to pennsylvanica by the 
ambiguous term of “variety, it is clear from the original description that he was ap- 
plying the name to an infrasubspecific form. In his check list published the next year, 
Smith (1903, Check List of the Lepidoptera of Boreal America, Amer. Entomol. Soc., 
Philadelphia, p. 61) listed it as “b pennsylvanica Sm.,” as though it were a subspecies, 
but again, it is clear from the general context that he did not differentiate between 
infrasubspecific forms and subspecies. Other authors of that period, such as Dyar, also 
failed to make this distinction or did so inconsistently. Barnes & McDunnough (1917, 
Check List of the Lepidoptera of Boreal America, Herald Press, Decatur, Illinois, p. 
96) did consistently make the distinction and treated pennsylvanica as a subspecies. 
The elevation of pennsylvanica to the rank of a species group name by Barmes & 
McDunnough far antedates that of Werny, and it appears that they should be cited as 
the authors in the event of its continued use in a specific sense. However, in the new 
Check List of the Lepidoptera of America North of Mexico (Hodges et al., in press) I 
have retumed expultrix and pennsylvanica to their former status as synonymic names 
based on forms, with their original authorship, and have had to add pennsylvanica 
Werny, 1966, to the synonymy of E. pudens. 

Werny’s 1966 work is not easily obtained, and the reader may wish to note that it 
was reviewed in this journal by J. C. E. Riotte (1969, J. Lepid. Soc. 23:101). 


DOUGLAS C. FERGUSON, Systematic Entomology Laboratory, IIBUI, Agricultural 
Research Service, U.S.D.A., % U.S. National Museum of Natural History, Washington, 
DEC» 20560) 


Journal of the Lepidopterists’ Society 
37(2), 1983, 181 


THE PROPER CITATION FOR THREE SPECIES NAMES 
PROPOSED BY S. H. SCUDDER 


The three names treated in this note are Chrysophanus dione, Apatura proserpina 
and Hesperia iowa. The three were each proposed in two different articles published 
sufficiently close to each other to cause some trouble. One of these articles is a report 
of the minutes of the meeting of the Entomological Section of the Boston Society of 
Natural History for 26 February 1868 in the Proceedings of the Society, vol. 11, p. 401, 
1868. The other article is “A preliminary list of the butterflies of lowa.” It appeared in 
the Transactions of the Chicago Academy of Science, 1:326—337, “1867-1870.” 

The following is the treatment of these names by cataloguers and checklisters from 
Kirby in 1871 to Miller & Brown in 1981. In it “B” stands for reference to the Boston 
publication and “C” to the Chicago one. The dates presented are abbreviated thus: ’68 
for 1868 and so forth. 


Catalogue or list Date C. dione A. proserpina H. iowa 
Kirby 1871 B ’68 B ’68 C ’69 
Strecker 1876 B 68 B ’68 C 68 
Edwards Sia C no date C no date B no date 
Edwards 1884 C ’69 € 69 C ’68 
Skinner 1896 C 69 B ’68 C ’68 
Dyar 1902 C ’69 B ’68 no data 
dosPassos 1964 68 68 68 
Miller & Brown 1981 C ’69 CaO C 69 


Barnes & McDunnough (1917), Bames & Benjamin (1926), and McDunnough (1938) 
cite no dates nor original description data; dosPassos cites dates but nothing more. 

Notice that only dosPassos is consistent. He at least cites the same year, 1868, for 
publication of the three names. Strecker also is consistent so far as the date is concerned 
but not for the publication in which the names were first published. Only W. H. Ed- 
wards, 1884, and Miller & Brown, 1981, are consistent for the periodical but not for 
the dates. What is the correct date and periodical? 

Publication of the names in the Proceedings of the Boston Society of Natural History 
was not directly by Scudder but by the secretary of the Entomological Section. This 
situation is covered by Article 50 (a) of the International Code of Zoological Nomen- 
clature. The names are to be credited to Scudder, not to Mann. Volume 11 was com- 
pleted and published in 1868. Scudder’s names became valid sometime after 26 Feb- 
ruary and before 31 December of that year. 

The only volume of the Transactions of the Chicago Academy of Science was pub- 
lished before 31 December 1870. Scudder dates his manuscript “Boston, October 22nd, 
1869.” Since none of the signatures of this journal are dated, publication must be 
considered “31 December 1870” according to the “Code.” The date on Scudder’s 
manuscript itself places the Chicago appearance of the names after their appearance 
in the Boston journal. 

The proper citation of each of the three names: Chrysophanus dione, Apatura pro- 
sperpina and Hesperia iowa, all sired by S. H. Scudder, is: Proceedings of the Boston 
Society of Natural History, 11:401, 1868. The signature containing the names is 26 
and is dated May 1868. 


F. M. Brown, Wright-Ingraham Institute, Colorado Springs, Colorado 80904. 


Journal of the Lepidopterists’ Society 
37(2), 1983, 182-186 


LIRIMIRIS MERIDIONALIS (SCHAUS), A NOTODONTID MOTH 
ASSOCIATED WITH COCOA (THEOBROMA CACAO L.) IN BELIZE 


Knowledge of the biology of the large Neotropical notodontid moth, Lirimiris meridi- 
onalis (Schaus), is limited to the early descriptions of adults of this species and allied 
ones from Costa Rica (Schaus, 1901, Trans. Entomol. Soc. London 1901:257-343; 1904, 
Trans. Amer. Entomol. Soc. 30:135-178; 1911, Ann. Mag. Nat. Hist. 7:262-285; 1912, 
Ann. Mag. Nat. Hist. 8:34-57). But, one early report from Argentina mentions that the 
caterpillar of the related species, L. lignitecta Walker, feeds on Chorisia insignis Kth. 
(Bombacaceae) and pupates in the soil (Schreiter, 1943, Acta Zool. Lilloana 1:7-44). 
Seitz (1907, Macrolepidoptera of the World, American Rhopalocera, Notodontidae 6: 
901-1452, A. Kernan, Stuttgart) was, therefore, correct in stating that, although the 
Notodontidae of the Neotropical Region were diverse, very little is known about the 
life cycles and larval food plant associations of most genera and species. This note 
reports the discovery of L. meridionalis feeding on cocoa, Theobroma cacao L. (Ster- 
culiaceae), in Belize. It constitutes not only the first published record of a larval food 
plant for this species but also the first description of the larval and pupal stages, in- 
cluding notes on behavior. 

During August 1981 I discovered three caterpillars of L. meridionalis feeding on the: 
mature leaves of T. cacao in Field Block 18 of the Hummingbird Hershey Cocoa Farm 
(88°38'W, 17°8’N) located about 18 road miles southeast of Belmopan. A good general 
description of this cocoa farm is available (Harler, Agribus. Worldwide, April/May 1981: 
22-31). The discovery of the caterpillars was made during the rainy season. They were 
found on a cocoa tree with a well developed leafy canopy. The caterpillars were col- 
lected and confined with fresh cuttings of cocoa leaves to a large, clear plastic bag for 
further rearing. At the time of collection, a ceratopogonid midge (Diptera) was found 
attached to the cuticle of one caterpillar. The midge, together with the caterpillar, was 
gently placed in a separate rearing bag for further observation, as the midge appeared 
to be feeding on the caterpillar. 

When collected the caterpillars were about 30 mm long, but they attained nearly 100 
mm in length (and 15-17 mm in width) by the time of pupation (3 September 1981 for 
the first one), following about three weeks in captivity. Each caterpillar molted once 
in captivity, suggesting that they were in the fourth instar at the time of discovery 
(assuming five instars prior to pupation). 

A macro-description of the caterpillar follows: Head capsule glossy butterscotch- 
yellow with two dorsal pairs of black dots, a central dot on each side, and three latero- 
ventral pairs. Background body color white with large black splotches. “Collar” con- 
necting head capsule with rest of body, yellow. First thoracic segment with dorsal 
black patch with white in center, lateral black mottling and a row of long, lateral white 
hairs. Second thoracic segment entirely white with one pair of dorsolateral white hairs, 
shorter than those of first segment. Third thoracic segment white with pairs of black 
spots, one central and two larger ones dorsolateral. Dorsolateral hairs also present. 
Segments 4 through 6 similar to 3, but segments 6 and 7 without hairs. Segment 6 
entirely white dorsally with a lateral black wedge-shaped streak continuous with in- 
tricate dorsal black pattern on 7. Segments 8 and 9 entirely white with small lateral 
black spot on 9. Conspicuous black wedge-shaped pattern on segments 10 through 12. 
Thin, irregular black line delineates lateral from ventral sections lengthwise; all legs 
and ventral cuticle butterscotch-yellow speckled with many small black dots. Last three 
segments enlarged, almost bulbous and thicker than the head capsule. These segments 
yellow with black speckling. Segments 9 through 11 with lateral long white hairs. 
Dorsolateral protuberances on last three segments with long white hairs, and a latero- 
ventral set on last segment as well. Anal plate shiny black. Together with anal plate, 
last three segments resemble a false head. When feeding, the caterpillar emits a loud 


VOLUME 37, NUMBER 2 183 


Fic. 1. Life cycle of the notodontid moth Lirimiris meridionalis (Schaus). Coun- 
terclockwise, top to bottom: biting midge Forcipomyia (Microhelea) fuliginosa (Mei- 
gen) (Diptera: Ceratopogonidae) feeding on the caterpillar of L. meridionalis at Hum- 
mingbird Hershey Cocoa Farm in Belize; fourth instar feeding on mature cocoa leaf; 
two cocoons with top cocoa leaf peeled away to show the flattened, compact structure; 
newly-eclosed adult moth. 


clicking sound, clearly audible from several feet away. During the rearing period, the 
appearance of the caterpillar did not change. But, it becomes an active prepupa, turning 
completely yellow-orange and the body contracts greatly in size. In captivity, the cat- 
erpillar makes a thin, dark-brown papery cocoon wedged between dead cocoa leaves; 


184 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Fic. 2. Below: pupal case, cast-off exuvium, and cocoon of L. meridionalis. Above: 
mounted specimen reared rom the Belize sennelle of caterpillars and deposited in the 
permanent collections of the Milwaukee Public Museum. 


VOLUME 37, NUMBER 2 185 


\) 


AA 


Fic. 3. Schematic line drawing of the fifth instar caterpillar of L. meridionalis. 


it is conspicuously flattened dorsoventrally. The life stages studied are shown in Figs. 
1 and 2, and an approximate schematic representation of the caterpillar’s appearance 
is given in Fig. 3. The cocoon is loosely made of coarse strands of silk holding leaves 
together to comprise a flat envelope. The pupal stage lasts 18 days under the rearing 
conditions employed. 

While either feeding or resting, the caterpillar assumes a tight “J” position, with the 
head curled around and partly concealed with the bulbous hind region. It was deter- 
mined that a biting midge, Forcipomyia (Microhelea) fuliginosa (Meigen) (Diptera: 
Ceratopogonidae) was sucking haemolymph from one caterpillar (Fig. 1). The discovery 
was made at 0900 h, and the midge remained in the position even when the caterpillar 
was transferred to a plastic bag from the tree. At 1700 h the same day, the midge was 
still attached at the same spot on the cuticle. By this time the caterpillar appeared 
traumatized, with the cuticle darkening. By 0700 h the following day the midge was 
no longer feeding and the caterpillar appeared healthy once again. This caterpillar 
survived to pupate and produce an adult. 

Although three caterpillars were found on the same cocoa tree, it is unlikely that L. 
meridionalis oviposits in clusters, as noted for some Temperate Zone notodontids (e.g., 
Farris & Appleby, 1980, J. Lepid. Soc. 34:368-371; Holland, 1916, The Moth Book, 
Doubleday, Page & Co., New York and Garden City, 479 pp.). Lirimiris meridionalis 
was originally described from adults collected in British Guiana (Schaus, 1904, op. cit.), 
and because the caterpillars, as for most notodontids with known life cycles, are tree- 
feeders, little else was discovered about this species. It is very doubtful that this species 
has been studied (R. W. Poole, pers. comm.). In the Temperate Zone notodontids have 
diversified considerably at the generic level in terms of larval food plant selection (e.g., 
McFarland, 1975, J. Lepid. Soc. 29:112—125; 1979, J. Lepid. Soc. 22(Suppl. 3):72 pp.), 
but sometimes a single species exhibits considerable polyphagy (Dirks, 1937, Maine 
Agric. Expt. Sta., Orono, Bull. 389, 162 pp.). In some tropical and subtropical regions, 
there is considerable diversification of larval food plant patterns among genera (e.g., 
Pinhey, 1975, Moths of Southern Africa, Tafelberg Publ., Cape Town, 273 pp.). Also 
noted to vary greatly among genera and species is the type of cocoon construction or 
pupation habit (Kendall, 1974, J. Lepid. Soc. 28:243-245; Farris & Appleby, op. cit.). 

There are no published records of L. meridionalis being associated with cocoa as a 
larval food plant (Costa Lima, 1936, Terceiro Catalogo Nos Insectos Que Vivem Nas 
Plantas Do Brasil, Minist. Agricult., Rio de Janeiro, 460 pp.; Entwistle, 1972, Pests of 
Cocoa, Longmans, London, 779 pp.). The association may be of economic interest since 
there are some records of other notodontids in the tropics being serious defoliators of 
fruit trees (e.g., Fujii & Yoshida, 1981, Proc. Hawaiian Entomol. Soc. 23:345-350), 
although some studies reveal very low densities of a single herbivorous insect species 
in large stands of a single food plant species (e.g., Solomon, 1981, Ecology 62:1205— 
1214). Although some Lepidoptera associated with cocoa in Brazil have tachinid (Dip- 


186 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


tera) parasites associated with them (Silva, 1980, Rev. Theobroma 10:257-259), the 
impact of biting midges such as F. (M.) fuliginosa on caterpillars is probably slight. 
Neotropical biting midges of the genus Forcipomyia and the subgenus Microhelea are 
ecto-parasites on a variety of plant-associated insects (Wirth, 1971, Entomol. News 82: 
229-245; 1972, Ann. Entomol. Soc. Amer. 65:564—577), and particularly the soft-bodied 
oairenoflllens of Lepidoptera (Wirth, 1972, J. Lepid. Soc. 26:65). 

The general lack of information on the association of L. meridionalis with cocoa 
anywhere in the Neotropical Region tentatively suggests that the interaction is very 
patchy regionally, even though this notodontid and related species are known from 
various localities in Central and South America. But, given the close evolutionary 
affinity between the Sterculiaceae and the Bombacaceae (Cronquist, 1981, An Inte- 
grated System of Classification of Flowering Plants, Columbia, New York, 1262 pp.), 
it is not surprising to find closely related species of Lirimiris associated with both 
tropical plant families (this report and Schreiter, op. cit.). Because of intense commer- 
cial interest in cocoa, there exists an unusually long list of herbivorous insects associat- 
ed with this fruit tree (e.g., Entwistle, op. cit.); but there is still a dearth of biological 
data on L. meridionalis. A fourth-instar caterpillar of L. meridionalis was discovered 
on T. cacao at Finca la Tigra, near La Virgen (10°23'N, 84°07’W), Heredia Province, 
Costa Rica, on 2 March 1983. The association of this moth with other Sterculiaceae 
warrants investigation. 

I thank Dr. R. W. Poole for confirming the determination of the moth, and Mss. Joan 
Jass and Susan Borkin with literature searches. Dr. Willis W. Wirth identified the biting 
midge. Christine Deniger prepared the line drawing in Fig. 3. Special thanks to Gordon 
R. Patterson of Hershey Foods Corporation for arranging my stay at Hummingbird 
Hershey, and to Norris Wade and his staff there for assistance and fine hospitality. This 
report is a by-product of research on cocoa funded by the American Cocoa Research 
Institute. 


ALLEN M. YOUNG, Invertebrate Zoology Section, Milwaukee Public Museum, Mil- 
waukee, Wisconsin 53233. 


Journal of the Lepidopterists’ Society 
37(2), 1983, 187-188 


BOOK REVIEW 


THE AUDUBON SOCIETY FIELD GUIDE TO NORTH AMERICAN BUTTERFLIES, by Robert 
Michael Pyle. 1981. Alfred A. Knopf, Inc., New York. Format 3%” x 7!” (approximate- 
ly). 917 pp., 759 color figures, pictorial keys, several halftone figures integrated into 
the text. Durable flexible binding. Price: $11.95. 


This book is one in a series of practical nature guides being published in standard 
format by the Audubon Society. They are designed for field use and appear to be rugged 
enough to withstand a fair amount of abuse. The size is convenient to fit into a day 
pack or large pocket. The flexible plastic cover should wear well. 

In reviewing a book of this nature one must consider the potential audience. The 
Introduction states: “This book is designed for everyone who wants to know how to 
identify butterflies in backyards, parks and gardens, as well as in woods and fields.” 
The book certainly meets this goal. It is not directed toward the serious taxonomist but 
should be of interest even to specialists. 

The book begins with the usual discussion of butterfly anatomy and biology accom- 
panied by appropriate illustrations. This section is followed by comments on survival, 
habitat, observing butterflies, methods of identification and nomenclature. A detailed 
section then describes how to use the guide. The Audubon Society has adopted a 
standard format for its nature guides that involves color, pattern, silhouette, and ge- 
ography. Three illustrative examples are provided to aid the reader in learning how to 
use the guide to identify an observed butterfly. The illustrations group butterflies by 
color and silhouette, and not according to taxonomic placement. Each double page of 
illustrations has a thumbtab guide in the mid-left margin denoting color, pattern, or 
silhouette. This enables the reader to locate quickly the section containing an unknown 
species. This method is standard throughout the guide series. 

The colored illustrations follow immediately the prefatory material and include some 
photographs of eggs, larvae, and pupae, as well as adults. Nearly all of the North 
American butterflies are shown in natural color; a few appear in the subsequent text 
in black-and-white. 

Most of the photographs are of naturally posed butterflies in the field, representing 
the specimen as it would be observed at rest. A few have been rotated in position, 
apparently for artistic purposes. Some of the photographs represent obviously pinned 
material (one specimen lacks its antennae), and some appear to be of either stunned 
or chilled specimens that have been posed on various substrates. For the most part the 
quality of the color reproduction is excellent, having been done by the Swiss firm of 
Nievergelt Repro AG, Zurich. A few obvious exceptions regarding color fidelity are 
Figs. 97, 98, and 116 (all sulphurs), and Fig. 265 (the Florida duskywing), which ex- 
hibits an unnaturally intensified iridescence. 

The focus in the illustrations is generally sharp and clear, with Figs. 117 and 635 
(the sleepy orange and astarte fritillary respectively) notable exceptions. The hesper- 
ine skipper photographs are generally superb, although a few are poorly illuminated 
and would present identification problems for the casual observer. The hairstreak pho- 
tographs are the best that I have ever seen, especially those of the Callophrys-Mitoura- 
Incisalia group. The color fidelity is very good. Despite the excellent photography, it 
_would be difficult to make positive field identification of many species without actually 
catching them, simply because they are too wary for prolonged observation. 

Common names are used in the color figure captions, and the reader is referred to 
the scientific text, which follows the plates, for further discussion. The technical section 
is arranged according to family in the order: Papilionidae, Pieridae, Lycaenidae, Rio- 
dinidae, Libytheidae, Nymphalidae, Satyridae, Danaidae, Hesperiidae, Megathymi- 
dae. A glossary and index complete the book. 

The scientific nomenclature follows that in ““A Catalogue/Checklist of the Butterflies 
of America North of Mexico” (Memoir No. 2 of the Lepidopterists’ Society). There are 
some radical changes in nomenclature from the 1964 dos Passos Checklist (Memoir 


188 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


No. 1 of the Lepidopterists’ Society). For Holarctic genera, names have been brought 
into agreement with European usage. Various subgeneric names have been elevated 
to full generic status. 

Quite a few new common names have been validated in the book, which is unfor- 
tunate. In 1980 a joint committee composed of members from the Lepidopterists’ So- 
ciety and the Xerces Society was formed to develop an approved list of common names. 
The committee report and recommendations have yet to appear. Consequently, further 
common name confusion may occur. In most cases the new vernacular names are 
descriptive of either larval host plant preference or adult maculation. 

In the technical section approximately one page is devoted to each species. The 
butterfly is first described and then compared with similar species, followed by infor- 
mation about life cycle, flight period, habitat and range. Notes on local habitats are 
included when pertinent, such as: “Huckleberry heaths in the higher parts of Yellow- 
stone Park are good places to seek clear examples of this sulphur’; with reference to 
Colias pelidne. Discussion is restricted to the species level; there is no discussion of 
subspecies, although color and pattern variations are mentioned. No authority names 
or dates are placed following the scientific names. This appears to be the style adopted 
for the Audubon guides, and not an omission on the part of the author. 

The book appears to be accurate and current. Even the recently described Clossiana 
acrocnema (Gall & Sperling) is included. The extinct Xerces blue is also illustrated. 
One omission was noted in the description of Epidemia dorcas, page 523. The book 
states that the life cycle is undescribed, when in fact, W. W. Newcomb published the 
life history in 1911 in the Canadian Entomologist. The butterfly shown in Fig. 396 as 
the hickory hairstreak appears actually to be one of the forms of the banded hairstreak 
(Fig. 394). These two species are difficult to separate. I would question combining two 
taxa that have been formerly called Agriades aquilo and A. glandon in North America 
into the single taxon A. franklinii (Curtis). Since the North American counterparts of 
aquilo and glandon exhibit the same habitat and host plant preferences as their Eu- 
ropean congeners, it would appear appropriate to use franklinii for the coastal races 
and rustica W. H. Edwards for the montane species. This comment, however, actually 
relates to the Catalogue, since it dictated the nomenclature used by Pyle. Lycaeua cu- 
preus is incorrectly listed as Chalceria cupreus. Considering the scope of this book, 
the points mentioned above are minor. 

This new Audubon Society guide should prove an invaluable aid to nature lovers 
and beginning lepidopterists. Even the serious amateur and seasoned specialist should 
find it a handy reference for identifying species from unfamiliar regions of North Amer- 
ica. In general the text is concise and the color photography excellent. It is unfortunate 
that not all species are shown in color. Within the format constraints placed by the 
guide series and the intended audience, Robert Pyle has done an admirable job. At 
the price of $11.95 the book is a real bargain and a highly recommended addition to 
any lepidopterist’s library. 


CLIFFORD D. FERRIS, Bioengineering Program, University of Wyoming, Laramie, 
Wyoming 82071. 


Journal of the Lepidopterists’ Society 
37(2), 1983, 189-192 


BOOK REVIEW 


BUTTERFLIES AND MOTHS OF NEWFOUNDLAND AND LABRADOR, THE MACROLEPI- 
DOPTERA by Ray F. Morris. 1980. Agriculture Canada, Research Branch, Publication 
1691. 407 pp., 40 text figs., 34 col. pls. Obtainable from Canadian Government Pub- 
lishing Centre, Supply and Services Canada, Hull, Quebec K1A OS9. $US 18.00, $CAN 
15.00. 


This attractively bound and well illustrated volume is a treatment of about 55 species 
of butterflies and 488 moths reported from Newfoundland and Labrador, giving sci- 
entific and common names, distribution, flight period, information on immature stages, 
34 colored plates showing almost every species, about 30 distribution maps, a check 
list, and a glossary of terms. The introduction consists of a short history of lepidopter- 
ological studies in the region and of sections on geography and climate, basic anatomy, 
development, and collecting of Lepidoptera. Keys and descriptions for identification 
are not included and mostly are not needed, as colored illustrations usually serve this 
purpose well for the larger Lepidoptera. The book deserves recognition as the first 
fully color-illustrated guide intended to cover all macrolepidoptera occurring in any 
state or province of North America. For the illustrations alone it is a bargain that no 
one interested in Canadian macrolepidoptera will want to miss. The quality of the 
plates is variable but mostly good; the quality of many of the specimens used for the 
photography could have been better. Typography is excellent, and there are almost no 
printing errors except a disconcerting omission of commas from numbers of four or 
more digits (e.g., 103 600 km? on page 16). I noted only one misspelled name; “paralis” 
should read parilis (p. 174), and one lapsus in a plant name: Viburnum in error for 
Vaccinium (p. 92). 

Of particular interest are records of two European noctuids reported from North 
America for the first time. These are Agrochola lota (Clerck) and Acronicta auricoma 
(F.). 

The book does suffer from a variety of shortcomings that should be discussed in some 
detail because of its potential biogeographic importance in documenting the fauna of 
one of the more interesting areas in North America. Morris does not seem to appreciate 
the geographical nature of subspecies, because in at least six instances he reports the 
occurrence of different subspecies of the same species in Newfoundland. All the Pa- 
pilio glaucus would surely have to be subspecies canadensis. The black female re- 
ported from Newfoundland by Clark & Clark, cited by Morris, is in the U.S. National 
Museum. It is very old (Oberthur collection) and probably mislabelled. The only ringlet 
in Newfoundland is Coenonympha tullia mcisaaci (the two specimens as figured are 
female and male of mcisaaci), and all Callophrys augustinus must surely be subspecies 
helenae. Similarly, one would think that there should be only one subspecies of Nym- 
phalis milberti, Carsia sororiata, and Dysstroma hersiliata. However, N. milberti viola 
is said to be concentrated mainly in the southern part of the island and nominate 
milberti to be more prevalent northward. If this observation is correct, then nominate 
milberti may be moving in from the mainland through Labrador and diluting the en- 
demic subspecies viola. Nominate Anomogyna perquiritata is reported from Labrador 
and subspecies beddeci from Newfoundland. But perquiritata was described from the 
White Mountains, New Hampshire, and I cannot see that New Hampshire material 
differs from beddeci in any significant way. I regard the latter as a junior synonym. 

I noted the following misstatements. Under Oeneis jutta (p. 45), how could Moschler 
have reported terraenovae from Labrador in 1860 when this subspecies was not de- 
scribed until 1935? The species of Vanessa (p. 58) do not hibernate as adults or pupae 
but are annual immigrants from much farther south, probably not overwintering any- 
where in Canada. The currently accepted family name for Ctenucha is Ctenuchidae 
rather than Amatidae (p. 78). Morris (p. 138) refers to my account of Leucania comma 
in Newfoundland as unpublished when indeed it was published (1963, Can. Entomol. 
95: 105-107). 


Considering that about 542 species are involved, errors of identification are few. 


190 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


However, identification problems of several kinds have found their way into the work. 
Sometimes the determinations of previous authors are accepted without question. Thus 
the records of Boloria chariclea are almost certainly based on misidentifications of B. 
titania. The former is a high arctic tundra species that would not be expected in New- 
foundland. Anomogyna fabulosa is treated as two species, being included both under 
its correct name and as A. sincera. The latter is a palearctic species with which the 
nearctic fabulosa was confused before being described as distinct in 1965. The record 
of Schrankia turfosalis is based on a misidentification of one of the similar North 
American species, and the correct generic name for this group is Hypenodes Double- 
day. Three species of Hypenodes, H. fractilinea (Sm.), H. palustris Fgn., and H. som- 
brus Fgn., are known from Newfoundland, but none is listed. Alypia octomaculata 
should have been deleted from the list or verified, as it is almost certain that the original 
record was based on misidentified specimens of A. langtoni. Confusion of these two 
species in the north where both might occur has become almost a tradition, but it 
should be noted that the hosts of octomaculata (Vitaceae) are absent from Newfound- 
land. Similarly, it would have been better to have disposed of Utetheisa idae in the 
introduction or a footnote instead of giving it formal listing as though it were really a 
part of the fauna. As Morris tells us, U. idae was described from Swain’s Island, New- 
foundland, in error for Swain’s Island, Samoa. 

Other misidentifications are simple errors of the author or of those who did identi- 
fications for him. The following should be noted: Polia leomegra, described from New- 
foundland, and P. carbonifera, from Alberta, refer to forms of the same species that 
should have been listed as leomegra. As it turns out, however, both are now regarded 
as synonyms of P. rogenhoferi, and the rogenhoferi that he lists was described in 1980 
as a new species, P. propodea McCabe, too late for inclusion in Morris’s book. Hy- 
phantria cunea and H. textor are generally regarded as one species, although Morris 
lists both. One is left guessing as to how he distinguished them. His figure of cunea 
on plate 10 is the immaculate form that has been regarded as textor, and his figure of 
textor appears to be Spilosoma congrua, not otherwise known from Newfoundland but 
possibly present. On plate 26 the figures of Malacosoma americanum (Figs. 1, 2) and 
M. disstria (Figs. 3, 4) are reversed. Pl. 28, Fig. 9, shows a specimen of Dysstroma 
truncata (not listed) as D. walkerata, although Fig. 10 is correctly determined as walk- 
erata. The latter is the peculiar black and white subspecies that occurs there. The 
species illustrated as Thera contractata (PI. 28, Fig. 16) is T. juniperata (L.), an intro- 
duced palearctic species now widely distributed in the Northeast. The report of T. 
otisi is puzzling because the illustration (Pl. 28, Fig. 17) really does look more like that 
species than like contractata. Otherwise, I would dismiss it as a probable misidenti- 
fication of contractata, which I have collected in Newfoundland myself. Pl. 31, Fig. 
6, shows an aberrant specimen that I would not recognize as Anacamptodes vellivolata, 
although it may be one. PI. 31, Fig. 14, appears to show a specimen of Homochlodes 
lactispargaria, not H. fritillaria as stated. Fig. 15 on the same plate shows the summer 
form of Plagodis phlogosaria, which would not be expected to occur in Newfoundland 
where there is no second brood. Only the spring form shown in Fig. 16 should be 
present. Differentiation between nominate Metarranthis duaria and its supposed 
northern subspecies, septentrionaria, is unsatisfactory because both were described 
from Canada. Whatever name is used, the Newfoundland population is certainly of the 
usual northern type and variable, as the species is everywhere. The U.S. National 
Collection has Newfoundland specimens even darker than that shown on PI. 32, Fig. 
2, the “subspecies” said not to occur there. The specimen shown on PI. 29, Fig. 25, as 
Perizoma basaliata is not that species but P. grandis Hulst. Both species occur in 
Newfoundland. Pl. 29, Fig. 32, shows what appears to be a specimen of Hydrelia 
condensata (Gn.) rather than inornata, the latter name being a synonym of lucata 
(correctly identified in Fig. 31). Inasmuch as Cerastis tenebrifera.does not occur in 
Nova Scotia, I question the recofds from Newfoundland and think it more likely that 
they were based on misidentified specimens of the closely similar Metalepsis fishi. 
The species of Hyppa reported as indistincta appears to be what I have identified 
from Newfoundland as H. brunneicrista Sm.; at least it almost exactly matches material 


VOLUME 37, NUMBER 2 191 


of the latter species from Alberta. The type of indistincta in the U.S. National Museum 
is something different. Pl. 33, Fig. 3, shows a specimen of Estigmene acrea arizonensis 
Roths. that must have come from the western U.S. In choosing an example for illustra- 
tion, the author apparently overlooked the fact that eastern males, including those from 
Newfoundland, always have yellow hindwings. 

The one most irritating feature of the book is its failure to indicate the geographical 
source of the illustrated specimens, especially those representing rare or doubtfully 
identified species. Obviously, some of those shown are from Newfoundland or Lab- 
rador, but many are not, and the permanent visual evidence that might have been 
afforded by the inclusion of label data in the legends is needlessly lost. Illustrations 
of the following species are among many for which specimen data would have been 
of considerable interest: Speyeria atlantis (does not look like subspecies canadensis); 
Spilosoma congrua (identified as Hyphantria textor); Arctia caja (not the arctic sub- 
species that occurs in Labrador); Agrotis volubilis (figure correctly identified as vol- 
ubilis, but is the specimen from Newfoundland? I had supposed, perhaps incorrectly, 
that A. musa replaces A. volubilis there); Agrotis obliqua (questionable record of a 
western species); Amathes c-nigrum (now Xestia spp.) (very pale hindwings; looks like 
a European specimen); Cerastis tenebrifera (questionable record); Cucullia asteroides 
(the figured specimen is this species, but its presence in Newfoundland is unlikely); 
Lithophane lepida (not the brightly marked form that one would expect in Newfound- 
land; looks like southern subspecies adipel Benj.); Trichoplexia exornata (figure does 
not appear to agree with the very large, distinctly marked form common in Newfound- 
land); Platysenta sutor (a southern species that occurs only as a casual immigrant 
northward); Epizeuxis aemula (appears to be the true aemula, although all Newfound- 
land material that I have seen belongs to a different, closely related species); Itame 
argillacearia (not in Nova Scotia); Itame exauspicata (not in Nova Scotia); Agrochola 
lota; and Acronicta auricoma (new North American records). 

Another criticism concerns the way in which life history information is cited. The 
statement, “Details of the immature stages in Newfoundland and Labrador are not 
available,” appears frequently, thus implying that such information, when given, is 
original or from some local source. Obviously, this is not so. I found no evidence in 
the introduction or elsewhere that any Lepidoptera were reared in connection with 
this project and concluded that the data were gleaned from many sources. I noted two 
conspicuous instances of misleading host information. Wax myrtle (Myrica cerifera) is 
an impossible host for any Rheumaptera species (p. 239) because it is a southern shrub 
that grows only where these moths do not occur. They do feed on Myrica gale and M. 
pensylvanica. In the discussion of Papilio brevicauda (pp. 33, 34), what has long been 
recognized as the major host plant is not mentioned. This is a seashore umbel, Ligus- 
ticum scothicum, whole stands of which sometimes may be decimated by larvae of this 
butterfly. The plants cited, Heracleum and Angelica, seem to be secondary hosts that 
are not much used where Ligusticum is available. 

Although Morris searched the literature extensively for Newfoundland and Labrador 
records, he overlooked a few. Psychophora phocata (Moschler) was described from 
Labrador. Hydriomena exculpata nanata McD. is represented from Hopedale, Lab- 
rador by a paratype in the Canadian National Collection (and there are specimens from 
Newfoundland in the British Museum (Nat. Hist.)). Covell (1970, Trans. Amer. Ento- 
mol. Soc. 96:145) reported Scopula limboundata (Haw.) from Grand Lake, Newfound- 
land. Anomogyna homogena conditoides Benj. was originally described from a large 
series from Salmonier, on the Avalon Peninsula. Forbes (1954, Cornell Exp. Stn. Mem. 
329:249) mentioned a specimen of an unidentified Merolonche species from Hopedale, 
Labrador. Although the location of that specimen is unknown to me, I verified the 
presence of such a species by collecting Merolonche ursina Sm. (otherwise a Rocky 
Mountain species) in southwest Newfoundland. In a paper on host records that I pub- 
lished in 1975 (U.S. Dept. Agric. Tech. Bull. 1521), several species are mentioned from 
Newfoundland, and two of these are not reported by Morris. The larva of Papaipema 
harrisi (Grt.) was collected from stems of cow parsnip, Heracleum lanatum, at Millville, 


192 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Codroy Valley, and Homochlodes lactispargaria (Wlk.) was reared on braken fern, 
Pteridium aquilinum, from a female collected at the same place. 


DOUGLAS C. FERGUSON, Systematic Entomology Laboratory, IIBIII, Agricultural 
Research Service, U.S.D.A., % U.S. National Museum of Natural History, Washington, 


D.C. 20560. 


Date of Issue (Vol. 37, No. 2): 6 September 1983 


EDITORIAL STAFF OF THE JOURNAL 
THOMAS D. EICHLIN, Editor 


% Insect Taxonomy Laboratory 
1220 N Street 
Sacramento, California 95814 U.S.A. 


MAGDA R. Papp, Editorial Assistant 


GLAS C. FERGUSON, Associate Editor THEODORE D. SARGENT, Associate Editor 


Continued from outside back cover 


. * The second specimen of Epargyreus spanna (Hesperiidae). Frank Gali & 


OSE GS SUN NOS Sc US uaa ML eA ale gC 170 

‘4 Battus zetides in the Republica Dominicana. Frank Gali & Albert 
| Se a AR AR SIR Sag RR Nl, os ano PO an Ae 171 

_ Pupal size and egg production characteristics in Rothschildia forbesi (Sa- 
meme mnnnonas As Miller 08s Oe) ee 174 

Occurrence of Megisto cymela (Satyridae) at flowers, with a behavioral 
MemCmEMnaNnRORTLIS!S. Williams). Ye a 176 

_ The “white male” variant of Colias (Pieridae): Two new records from Col- 
Ce TSE a2 i OT UI RIS Mer ate ptt 00 call Up eae Cae rede Senor Ua Ler: 

On the status of Pseudothyatira expultrix (Grt.) and Euthyatira pennsyl- 
vanica J. B. Smith (Thyatiridae). Douglas C. Ferguson 00.000 179 

The proper citation for three species names proposed by S. H. Scudder. F. 
CE SOE IAIN STC ASE USE TiO ROGCAE sl Dk RAY er NE eM Neale Ace eT Sort 181 

Lirimiris meridionalis (Schaus), a notodontid moth associated with cocoa 
(Theobroma cacao L.) in Belize. Aller M. YOur gy o.:esscos.e-ssseeseeeeevvneseereeeveeen 182 
EE SS Sr TE le ence a Ne ae ERA 167,189 


Ares, 


Jorrespondence: Address all matters relating to the Journal to the editor. Short 
nuscripts such as new state records, current events, and notices should be sent to the 
editor of the News: June Preston, 832 Sunset Drive, Lawrence, Kansas 66044 U‘S.A. 


PRINTED BY THE ALLEN PRESS, INC.. LAWRENCE, KANSAS 66044 U.S.A. 


CONTENTS 


NATURAL HISTORY OF SEVEN HAIRSTREAKS IN COASTAL NORTH 
CAROLINA. Samuel M. Gifford & Paul A. Opler 


NOTES ON THE SATYRID BUTTERFLY POPULATIONS OF COR- 
COVADO NATIONAL PARK, Costa Rica. Paul L. Whit- 
EGRET 6 LE Wy a SO 


A NEW SUBSPECIES OF SPEYERIA EGLEIS (NYMPHALIDAE) FROM 
THE PUMICE REGION OF CENTRAL OREGON. Paul C. 
Hammond & Ernst J. Dornfeld ae 


CAUSAL ANALYSIS OF A MIGRATION OF THE SNOUT BUTTERFLY, 
LIBYTHEANA BACHMANII LARVATA (STRECKER) (LIBY- 
THEIDAE). ‘Raymond W. Neck ... | 


LEPIDOPTERA ASSOCIATED WITH WESTERN SPRUCE BUDWORM 
IN THE SOUTHWESTERN UNITED STATES. Robert E. Ste- 
vens, V. M. Carolin ¢> Catherine Stein 


Two NEw SPECIES OF THE TRIBE EUCOSMINI (TORTRICIDAE) 
CLOSELY RELATED TO PHANETA  GRANULATANA 
(KEARFOTT). Andre Blanchard & Edward C. Knudson .... 


THE IDENTITY OF Two MONOTYPIC GEOMETRID GENERA 
WRONGLY ATTRIBUTED TO THE NEARCTIC FAUNA (GEO- 
METRIDAE).: Douglas C. Ferguson... 3 


A NEw SPECIES OF SHINIA (NOCTUIDAE) FROM CENTRAL FLORI- 
DA, WITH DESCRIPTION OF ITS LIFE History. D. F. 
Hardwick ic 


A NEw SPECIES OF EOMICHLA FROM CosTA RICA (OECOPHORI- 
DAE). J. F. Gates Clarke > 


A BRIEF DESCRIPTION OF THE PHYSIOLOGICAL TECHNIQUES— 
Disc GEL ELECTROPHORESIS, INCLUDING GEL PHO- 
TOGRAPHY AND THIN LAYER CHROMATOGRAPHY. 
Thomas BR. Taylor a 


GENERAL NOTES ) 
Overwintering aggregations of hackberry caterpillars (Asterocampa clyton: 
Nymphalidae)... Nancy E, Stamp 02 

Myscelia antholia (Nymphalidae) in the Reptblica Dominicana. Frank 
Galids Albert:Schwartz 0 

Notes on the autumnal northward migration of the cloudless sulphur, Phoe- 

bis sennae (Pieridae), along the South Carolina coast. L.L. Gaddy & 


Pete Laurie. ih oh 
Aberrant Hemileuca maia (Saturniidae).  Johmr V. Calor ieeeeceeesessceennneneeee 
Noctua pronuba (L.) on Sable Island, Nova Scotia, a record of dispers- 

al: Barry Wright i 300 


Continued on inside back cover 


_ Volume 87 1983 Number 3 
ISSN 0024-0966 


JOURNAL 


of the 


LEPIDOPTERISTS’ SOCIETY 


Published quarterly by THE LEPIDOPTERISTS’ SOCIETY 


. Publié par LA SOCIETE DES LEPIDOPTERISTES 
_ Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN 
Publicado por LA SOCIEDAD DE LOS LEPIDOPTERISTAS 


DEDICATED TO THE MEMORY OF 
Ernst J. Dornfeld 
1911-1983 


27 April 1984 


THE LEPIDOPTERISTS’ SOCIETY 


EXECUTIVE COUNCIL 


LEE D. MILLER, President CHARLES V. COVELL, JR., 
KAROLIS BAGDONAS, Vice President Immediate Past President 
MIGUEL R. GOMEZ BUSTILLO, Vice President JULIAN P. DONAHUE, Secretary 
J. DONALD LAFONTAINE, Vice President RONALD LEUSCHNER, Treasurer 


Members at large: 


K. S. BROWN, JR. F. S. CHEW J. M. BurNs 
E. D. CASHATT G. J. HARJES F. W. PRESTON 
T. C. EMMEL E. H. METZLER N. E. STAMP 


The object of the Lepidopterists’ Society, which was formed in May, 1947 and for- | 
mally constituted in December, 1950, is “to promote the science of lepidopterology in 
all its branches, .... to issue a periodical and other publications on Lepidoptera, to facil- 
itate the exchange of specimens and ideas by both the professional worker and the ~ 
amateur in the field; to secure cooperation in all measures’ directed towards these aims. 

Membership in the Society is open to all persons interested in the study of Lepi- 
doptera. All members receive the Journal and the News of the Lepidopterists Society. 
Institutions may subscribe to the Journal but may not become members. Prospective ~ 
members should send to the Treasurer full dues for the current year, together with their — 
full name, address, and special lepidopterological interests. In alternate years a list of 
members of the Society is issued, with addresses and special interests. There are four _ 


numbers in each volume of the Journal, scheduled for February, May, August and | 


November, and six numbers of the News each year. 


Active members—annual dues $18.00 
Student members—annual dues $12.00 
Sustaining members—annual dues $25.00 
Life members—single sum $250.00 
Institutional subscriptions—annual $25.00 


Send remittances, payable to The Lepidopterists’ Society, and address changes to: 
Ronald Leuschner, 1900 John St., Manhattan Beach, California 90266 U.S.A. 


Back issues of the Journal of the Lepidopterists’ Society, the Commemorative Vol- 
ume, and recent issues of the NEWS are available from the Treasurer. The Commem- 
orative Volume, is $6; for back issues, see the NEWS for prices or inquire to Treasurer. 


Order: Mail to Ronald Leuschner, 1900 ion St., Manhattan Beach, California 90266 ‘* 
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Journal of the Lepidopterists’ Society (ISSN 0024-0966) is published quarterly by the — 
Lepidopterists’ Society, a non-profit, scientific organization. The known office of publi- 
cation is 1041 New Hampshire St., Lawrence, Kansas 66044. Second class postage paid 
at Lawrence, Kansas, U.S.A. 66044. 


Cover illustration: Adult of the squash vine borer, Melittia cucurbitae (Harris) (Sesi- 


idae), which occurs in the eastern half of the United States and along the Gulf Coast into 
Vera Cruz, Mexico. The larvae are destructive borers in the vines of various cultivars of 
Cucurbita spp. (squash, pumpkins and gourds). Original drawing by Dr. Charles S. Papp, 
Sierra Graphics & Typography, 1722 J Street #19, Sacramento, CA 95814, USA. . 


JOURNAL OF 


Tur LEpPIDOPTERISTS’ SOCIETY 
Volume 37 1983 Number 3 


Journal of the Lepidopterists’ Society 
37(3), 1983, 193-206 


A NEW CLEARWING MOTH (SESIIDAE) FROM CENTRAL 
AMERICA: A STEM BORER IN MIMOSA PIGRA 


THOMAS D. EICHLIN 


Systematic Entomology Laboratory, Division of Plant Industry, 
Department of Food and Agriculture, Sacramento, California 95814 


AND 


STEVEN PASSOA 


Department of Entomology and Nematology, University of Florida, 
Gainesville, Florida 32611 


ABSTRACT. The eggs, last instar larva, pupa, and adult of a new species of Sesiidae, 
Carmenta mimosa, are described. This species is known to occur from southern Mexico 
to Nicaragua and has been reared from Mimosa pigra L. 


This species is being described now to provide a name for a publi- 
cation on insects injurious to Mimosa pigra L. (Leguminosae) in Hon- 
duras. This plant, native from Mexico to Argentina, is a serious weed 
pest in parts of northern Australia and Thailand. A survey to seek 
potential biological agents for control of the weed was conducted in 
1981. This complements work by Australian scientists looking for nat- 
ural enemies of M. pigra mainly in Brazil. 

Engelhardt (1946) included 24 North American species in his con- 
cept of the genus Carmenta. MacKay (1968) in her study of the larvae 
of North American species restricted the genus Carmenta by placing 
some species of the “complex” in her “new genus I” and in Alcathoe 
Hy. Edw., based solely on larval characters. Duckworth and Eichlin 
(1977a) considered 25 species to be in the genus north of Mexico, three 
more having been added to the list since that publication. Heppner 
and Duckworth (1981) list 31 species of Carmenta worldwide, mostly 
from the New World but a few from the Australian region. Preliminary 
sorting of the Neotropical species by Eichlin reveals that the genus will 
eventually contain many more species (200+) once this portion of 


194 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Fic. 1. Carmenta mimosa, n. sp., allotype female, near Linares, Nuevo Leon, Mex- 
ico. 


major revisionary studies on Western Hemisphere Sesiidae has been 
concluded (Duckworth & Eichlin, in prep.). 
One species from Argentina and Chile, C. haematica (Ureta), has 
ee recommended for biological control of broomweed, Gutierrezia 
p. (DeLoach, 1980). 


Carmenta mimosa Eichlin and Passoa, new species 


(Figs. 1-22) 


Description. Male: Head (Figs. 2-3) with vertex brown-black; front brownish gray, 
some white scales laterally; proboscis present, coiled; occipital fringe white or pale yellow 
dorsally, brown-black laterally with a few white scales mixed; antenna relatively short, 
thickened apically, less than one-half of forewing length, ciliate ventrally, dorsally brown- 
black with yellow powdered to apex, strongest apically; labial palpus slightly exceeding 
top of front, expanded somewhat laterally with roughened scales but sculptured ventrally 
appearing flattened, brown-black, strongly mixed with white ventrally, some pale yellow 
dorsally and on apical segment. Thorax brown-black, with narrow subdorsal yellow stripes; 
laterally with pale yellow at wing base and anteriorly. Abdomen constricted at base, 
brown-black, pale yellow on anterior half of segment 1, narrow pale yellow bands on 
posterior margin of segments 2, 4, 6 and 7 but may vary from bands only on 2 and 4 to 
bands on all segments, the latter condition apparently uncommon; ventrally, character- 
istically with segments 1 and 2 white, segment 4 pale yellow on posterior half, scales on 
posterior end forming keel; anal tuft not conspicuous at rest, narrowed to blunt point, 
laterally downwardly appressed, brown-black with some white scales mixed laterally. 


~ VOLUME 387, NUMBER 3 195 


Fics. 2, 3. Scanning electron micrographs of head of Carmenta mimosa, n. sp. 2 
(left), side view, x44; 3 (right), close-up of ocellus, <x 320. 


Legs mostly brown-black, pale yellow at base of forecoxa; some white mixed on tibial 
tufts, on spurs and powdered inside. Forewing mostly hyaline, margins very narrow, 
brown-black with pale orange to pale yellow powdered on discal spot, and margins, 
including apically between veins; more strongly powdered on underside of wings. 
Hindwing hyaline; costal margin powdered pale yellow; fringe concolorous with margins, 
becoming white at wing base. Wing length of male and female, 6-9 mm. Male genitalia 
(Figs. 4a, b) typical of Carmenta species, having elongate saccus approximately one-half 
length of valva, and saccular ridge apically abruptly downcurved to ventral margin of 
valva. 

Female (Fig. 1): Antenna as on male but lacking ventral ciliation. Maculation much 
like described for male, perhaps with broader apical margin on forewing. Female gen- 
italia (Fig. 5) typical for the genus, having ductus bursae pigmented and sclerotized for 
more than half its length (about two-thirds), with ductus seminalis originating from 
ductus bursae anteriorly beyond middle. 

Egg (Figs. 6, 7). Somewhat obovoid in shape, 0.50 + 0.03 x 0.83 + 0.02 mm, with 
anterior end (micropylar end) slightly flattened; posterior end rounded; top rounded, 
perhaps slightly depressed in center, bottom flattened. Surface of chorion minutely 
bumpy, resembling skin of orange; covered by aeropyles; reticulate pattern of low narrow 
ridges which intersect to form irregular, mostly hexagonal designs. Micropyle consisting 
of nearly triangular shaped pit with hole at each corner, surrounded by rosette of about 
seven oblong petal-like primary cells, which are then surrounded by approximately 11 
larger secondary cells of similar shape; surface within micropylar cells smoother than 
surface of other cells, with reduced number of aeropyles. 

Larva (Figs. 9-19). Body cream colored; head tan-brown; tonofibrillary platelets faintly 
visible, forming two wide horizontal bands; adfrontals nearly reaching vertical angle; six 
stemmata: stemmata 5 and 6 separated from 1-4; prothoracic shield with two oblique 
lines, one on each side of midline; below this is group of pigmented spots; pinacula 
inconspicuous, concolorous with body. Mouthparts: Mandible with four teeth; two lateral 
mandibular setae present; spinneret about two times length of basal segment of labial 
palps; hypopharyngeal complex with proximomedial region membranous, each side with 
long ridge covered by fine spines; two minute stipular setae present. Crochets: Homoi- 
deous, uniserial, fewer on A6 compared to A7 (x = 31:39 total number respectively); anal 
prolegs have small papillae above anal crochets. Chaetotaxy (setae as in MacKay, 1968): 


196 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Fics. 4,5. Genitalia of Carmenta mimosa, n. sp. (ventral view). 4a, b, male genitalia; 
3, female genitalia. 


VOLUME 387, NUMBER 3 197 


Fics. 6, 7. Scanning electron micrographs of egg of Carmenta mimosa, n. sp. 6 
(left), top view (micropyle on right end), x180; 7 (right), close-up of end with micro- 
pylar area, x 600. 


Head with Adfl and Adf2 widely separately from each other; frontal setae in line with 
frontal pores; clypeal setae as shown; labrum with medial and lateral setae as shown. 
Epicrania with three vertical setae in straight line angled toward Adf2; P1 long, below 
P2; L1 and A8 above Al and A2; Ol between stemmata 2 and 3; O2 below stemma 1; 


, pe A gee am 
os A MEE LP “ »eifler—" 


«lle ¢ ps 


Fic. 8. Larva of Carmenta mimosa in chamber exposed in branch of host plant, 
Mimosa pigra. 


198 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


12 13 


Fics. 9-18. Carmenta mimosa larva. 9, epicrania and front; 10, labrum; 11, cro- 
chets on abdominal segment one; 12, mandible, ventral view; 13, spinneret and hypo- 
phayrngeal complex. (See p. 206 for abbreviations used in figures 9-19.) 


VOLUME 37, NUMBER 3 199 


Fics. 14-16. Carmenta mimosa larval chaetotaxy (lateral view). 14, prothorax; 15, 
mesothorax; 16, first abdominal segment. 


SO3 below stemma 6; SO2 between stemmata 5 and 6; O3 remote from SO3 and stemma 
6. Prothorax with D2 longer than D1; XD1, XD2 and SD1 in a vertical row, SD2 smaller, 
behind and below XD2; prespiracular group trisetose, L3 much shorter than L1 or L2, 
all on triangular pinaculum; two SV setae widely spaced. Mesothorax with D2 longer 
than D1, both on same pinaculum; SD1 on same pinaculum with SD2; the position of 
L1 variable, more commonly close to L3, or equidistant between L3 and L2; one sub- 
ventral seta behind and below L2. Abdominal segment one with D2 below D1; SD2 
minute; SD1 above spiracle; L2 and L1 below spiracle on same pinaculum; L3 closer to 
SV3 than to L1; SV group bisetose, on same pinaculum. Abdominal segment seven with 
D2 longer than D1, each one on an oblong transverse pinaculum; SD1 above spiracle; 
SD2 minute, in front of spiracle; L1 and L2 on same pinaculum, below spiracle; L8 
widely spaced from L1; SV group bisetose, on same pinaculum. Abdominal segment 
eight with D2 above D1; SD1 below D2 and D1; L1 longer than L2; SD2 minute, in 
front of L2 + L1; L3 below SD2; one SV seta present. Abdominal segment nine with D2 
above D1 and SD1, latter two setae on one pinaculum; L2 present but reduced; L1 above 
SV1. Anal shield with SD1, L1 and D2 along margin; D1 smaller and set inward; one 
ventral seta present on all segments. 

Pupa (Figs. 20-23). Uniformly tan-brown in color. Head: Frons with projecting cir- 
cular ridge and two large carinae near eyes; labrum triangular, with 4 setae, inner ones 
largest; mandibles elevated; maxillary palps present, almost reaching maxillae; antennae 
about % length of wings; maxillae long, ending slightly before caudal margin of wings; 
labial palps present, widest in middle. Thorax: Prothoracic femur exposed; dorsum of 
prothorax with single median transverse ridge, mesothorax with one median ridge flanked 


200 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


SVI 17 


Fics. 17, 18. Carmenta mimosa larval chaetotaxy (lateral view). 17, seventh abdom- 
inal segment; 18, abdominal segments 8 and 9. 


on each side by two furrows; prothoracic leg about half length of maxillae; only tips of 
metathoracic legs exposed. Abdomen: A2 and A8-9 with single transverse row of stout 
spines dorsally; A3-6 with two rows of dorsolateral spines, those on anterior margin larger 
and more stout than those on posterior margin; on A7 males have two rows, females one; 
cremaster absent, in its place eight flattened spines irregularly spaced in circular pattern 
with their tips heavily sclerotized. 

Host. Mimosa pigra L. (Figs. 24-25). 

Distribution. MEXICO: Nuevo Leon, Michoacan and Oaxaca; HONDURAS and NIC- 
ARAGUA. 

Types. HOLOTYPE: 6, HONDURAS, 30 km SE Siguatepeque, VI-15-1979, J. A. Chem- 
sak, collector (California Insect Survey, Berkeley). ALLOTYPE: 2, MEXICO, Nuevo Leon, 
18 mi. W Linare, IX-17-76, J. A. Powell & J. A. Chemsak, collectors (California Insect 
Survey, Berkeley). PARATYPES 21: 266, 12, HONDURAS, Dept. Comayagua, Comayagua, 
reared ex. Mimosa pigra, S. Passoa, coll. I-9&10-81, (1 4, 1 2) emerged J-12-81, (1 4) 
emerged II-10-81, Genitalia Slide CDA #520, (1 2) Genitalia Slide CDA #610; 13 48, 
HONDURAS, Yoro, El Progreso, IV/10-15/79, C. Gentry, Pherocon 1C trap baited w 
Z,Z-ODDA, (1 8) Genitalia Slide CDA #522. 1 6, NICARAGUA, San Ramon, Matagalpa, 
VII-11-78, C. Gentry, Pherocon 1C trap baited w Z,Z-ODDA. 2 66, MEXICO, Oaxaca, 
Temascal, K. H. Janzen, collector (California Insect Survey, Berkeley), (1 6) X-10-1963, 
(1 6) X-3-1963, Genitalia Slide CDA #094; 1 9, MEXICO, Michoacan, Cotija, [X-14- 


VOLUME 387, NUMBER 3 


201 


LO A ees 
ag Se ns 

Cateye Zz 
Sig ples Sees. 

Se EN < 
Seal aS Z z 
Cat Pics Z 
CA pa eee << S 
< < =< 
Zoos < < 
OR = 3 
< S < 


23 


Fics. 19-23. Carmenta mimosa, larva and pupa. 19, anal shield, dorsal view; 20, 
pupa, ventral view; 21, close-up of pupal abdominal spines; 22, pupal abdomen, lateral 
view, male (female lacks the indicated row); 23, caudal view of pupal terminal abdom- 
inal segment. 


202 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Fics. 24, 25. Mimosa pigra, host plant for Carmenta mimosa, Comayagua, Hon- 
duras. 


VOLUME 37, NUMBER 3 203 


1975, B. Villegas (University of California, Davis). Paratypes will be distributed among 
the two California institution collections listed above; Florida State Collection of Arthro- 
pods, Gainesville; National Museum of Natural History, Washington, D.C., and Passoa’s 
collection. 

Also, the following material was studied: HONDURAS, Dept. of Comayagua, on Mi- 
mosa pigra (5 preserved larvae and 4 pupae associated with the reared adults mentioned 
above): 3 larvae in alcohol, 2 on slides with skin (lepidoptera mandible slides #200, 201, 
S. Passoa collection), 4 pupae, 2 cast pupal exuviae (one with larval skin mounted, 
lepidoptera mandible slide #19, Fla. State Coll. of Arthropods), 1 in alcohol (died close 
to eclosion, female genitalia dissected out, genitalia slide #267, S. Passoa Coll.), 1 pinned, 
Honduras, Dept. of Comayagua, Comayagua, XII-8-1981, in stem of Mimosa pigra, 
larva, not fully grown, lepidoptera mandible slide #18, Fla. State Coll. of Arthropods. 
This material will be placed in the Florida State Collection of Arthropods, University of 
California (Berkeley), and Passoa’s private collection. 


Discussion. With its clear wings, slender legs, shortened antennae, 
abdomen with narrow pale banding and constricted “waist,” labial 
palps sculptured to resemble mandibles when viewed face-on, C. mi- 
mosa effectively creates the impression of being a small species of 
vespid wasp. In Honduras the genus Polybia (Hymenoptera: Vespidae) 
resembles C. mimosa in markings, and this wasp is relatively common. 

As reported in the above data the adults from Comayagua, Honduras 
were reared from Mimosa pigra. A male and female emerged on 12 
January 1981 from pupae extracted from the branches a few days 
earlier. Another male emerged on 10 February 1981 from a section of 
branch where the larva was first observed on 9 January. Another larva 
was collected at the same clump of plants a year later (Fig. 8). 

The 13 male adults from E] Progreso, Honduras were collected from 
10 April to 138 June 1979 in sticky traps baited with a sex attractant 
containing about 96% of the Z,Z isomer of 3,13-octadecadien-1-ol ace- 
tate (Z,Z-ODDA), a major component of the sex pheromone systems 
of various sesiids (Duckworth & Eichlin, 1977b; Karandinos et al., 1977; 
Sharp et al., 1978; Sharp & Ejichlin, 1979; Neal & Eichlin, 1983; Niel- 
sen, 1979; Reed et al., 1981). An additional male was captured in the 
same manner in Matagalpa, Nicaragua, 11 July 1978. 

The Mexican specimens were captured in September and October. 
From the limited label data it appears as though the flight period 
extends over a long time (April to October); or the flight period varies 
depending on the locality; or there is more than one generation per 
year; or the actual situation involves some combination of the above 
possibilities. 

Carmenta prosopis (Hy. Edwards), a slightly smaller species on the 
average (wing length 5-8 mm) occurring from northern Mexico into 
southwestern United States, has been reared on several occasions from 
species of Prosopis (mesquite) and from Mimosa biuncifera Benth., 
where they developed as inquilines in hymenopteran-caused stem galls 


204 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


(Engelhardt, 1946). C. prosopis also has the somewhat unusually short- 
ened antennae found on C. mimosa but lacks the narrow waist and 
differs in several details of maculation and structures of the genitalia. 
Two other species in a closely related genus, Aegerina harti (Druce) 
and A. vignae (Busck, 1929), are known to be borers in various legu- 
minous plants, some of which are cultivated species (Duckworth & 
Eichlin, 1978). These moths also have shortened antennae and con- 
stricted abdomens but differ from C. mimosa mainly in details of 
genitalic morphology. 

In Mosher’s key (1916), the pupae of C. mimosa will key out with 
Podosesia Moschler and Memythrus Newman (=Paranthrene Hiib- 
ner), due to the unarmed clypeus. However, Paranthrene are larger, 
20-25 mm, than C. mimosa which measures only 8-15 mm. Podosesia, 
as illustrated by Mosher, differs from C. mimosa in that the maxillae 
extend ¥, of the way to the caudal margins of the wings. In C. mimosa 
they extend ¥, that distance. 

The larva of C. mimosa agrees with MacKay’s (1968) definition of 
the genus in having both D1 and D2 on large pinacula, especially on 
abdominal segment seven. The prespiracular group is arranged in an 
equilateral triangle rather than in an obtuse angle as in other members 
of the genus Carmenta as defined by MacKay. Specific identification 
will likely depend on small differences in the size and shape of the 
various pinacula. Unfortunately, these pinacula can be difficult to see 
unless the larval skin is cleared. Certain pinacula lacking setae are 
present but were not named by MacKay. In this study they are all 
illustrated but also left unnamed. 

External symptoms of larval damage include exit holes in the stems 
of the host and frass extruding from the holes. C. mimosa apparently 
was not seriously damaging its host in Honduras. However, larvae were 
collected from the same clump of plants in successive years, and such 
cumulative damage might weaken the plant in the long run. C. mi- 
mosa might be effective as a biological control agent in areas where 
strong winds and heavy rains make such a weakness in the stem a 
liability to the plant. Although the borer was never collected from any 
commercial crops in Honduras during a three year period (Passoa, 
unpub. data), some related species attack economically important le- 
gumes. Host specificity tests on this insect should include legumes. The 
use of pheromones for sampling populations could allow for easy eval- 
uation of establishment if releases are made. 


ACKNOWLEDGMENTS 


We thank the following individuals and their institutions for the loan of material used 
to describe this species: D. H. Habeck, University of Florida, Gainesville; J. A. Powell, 


VOLUME 87, NUMBER 3 205 


University of California, Berkeley; R. O. Schuster, University of California, Davis. We 
are grateful to C. R. Gentry, USDA/SEA-AR, Southeastern Fruit and Tree Nut Labo- 
ratory, Byron, Georgia, for trapping the series of specimens with attractants in Honduras 
and supplying them for this study. Thanks also to J. H. Tumlinson, USDA/SEA-AR, 
Insect Attractants, Behavior and Basic Biology Research Laboratory, Gainesville, Florida, 
for providing attractants used in this and most of our research. We appreciate the con- 
structive reviews of the manuscript by G. Buckingham and D. Habeck, University of 
Florida, Gainesville. Preparation of genitalia slides and other technical assistance were 
provided by Magda R. Papp, Biological Technician, California Department of Food and 
Agriculture, Sacramento. 

Special thanks are due to Kyra Carroll, Peace Corps (Honduras); her company during 
long field trips, sharp eyes for collecting, and help in choosing field sites are appreciated. 
The field work in Honduras was supported by the International Plant Protection Center, 
Aquatic Weed Program, at the University of Florida. We wish to thank M. Fonseca, 
Agriculture Director, Peace Corps, Honduras, for his guidance. G. Reyes, Regional Di- 
rector, Recursos Naturales, Comayagua, kindly provided office space and equipment 
needed to carry out this study. 


LITERATURE CITED 


Busck, A. 1929. A new aegeriid on cowpea from Brazil (Lepidoptera: Aegeriidae). 
Proc. Entomol. Soc. Washington 31:134-187. 

DELOACH, C. J. 1980. Prognosis for the biological control of weeds of southwestern 
United States rangelands. Pp. 175-199, in E. S. Del Fosse (ed.). Proc. Fifth Int. 
Symp. on Biological Control of Weeds. Brisbane, Australia. 

DUCKWORTH, W. D. & T. D. EICHLIN. 1977a. A classification of the Sesiidae of America 
north of Mexico (Lepidoptera: Sesioidea). Calif. Dept. of Food and Agric., Occas. 
Pap. Entomol., No. 26:1-54. 

1977b. Two new species of clearwing moths (Sesiidae) from eastern North Amer- 

ica clarified by sex pheromones. J. Lepid. Soc. 31:191-196. 

1978. The type-material of Central and South American clearwing moths (Lep- 
idoptera: Sesiidae). Smiths. Contrib. Zool. 261:1-28. 

ENGELHARDT, G. P. 1946. The North American clear-wing moths of the family Ae- 
geriidae. Smiths. Instit., U.S. Nat. Mus. Bull. 190:1-222. 

HEPPNER, J. B. & W. D. DUCKworTH. 1981. Classification of the Superfamily Sesioidea 
(Lepidoptera: Ditrysia). Smiths. Contrib. Zool. 314:1-144. 

KARANDINOS, M. G., J. H. TUMLINSON & T. D. EICHLIN. 1977. Field evidence of 
synergism and inhibition of the Sesiidae sex pheromone system. J. Chem. Ecol. 3: 
57-64. 

MacKay, M. R. 1968. The North American Aegeriidae: A revision based on last instar 
larvae. Mem. Can. Entomol. Soc. 58:1-112. 

MOsHER, E. 1916. A classification of the Lepidoptera based on characters of the pupa. 
Bull. Illinois State Lab. Nat. Hist. 12:13-159. 

NEAL, J. W., JR. & T. D. EICHLIN. 1983. Seasonal response of six male Sesiidae of 
woody ornamentals to clearwing borer (Lepidoptera: Sesiidae) lure. Environ. Ento- 
mol. 12:206-209. 

NIELSEN, D. G. 1979. Clearwing moth pheromone research: A perspective. USDA/ 
SEA, Agric. Res. Results 6:75-83. 

REED, D. K., T. D. EICHLIN & G. L. REED. 1981. Effectiveness of blends of synthetic 
sex attractants and comparisons with virgin female lesser peachtree borers as bait 
for capture of Sesiidae. Environ. Entomol. 10:488-—491. 

SHARP, J. L., J. R. MCLAUGHLIN, J. JAMES, T. D. EICHLIN & J. H. TUMLINSON. 1978. 
Seasonal occurrence of male Sesiidae in north central Florida determined with pher- 
omone trapping methods. Fla. Entomol. 61:245-250. 

SHARP, J. L. & T. D. EICHLIN. 1979. Distribution and seasonal occurrence of Sesiidae 
(Lepidoptera) attracted to E,Z and Z,Z acetate and alcohol. USDA/SEA, Agric. Res. 
Results 6:35—46. 


206 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


icine used in figures (9-19) of immature stages: 
(Al, A2, A3)—first, second and third anterior setae 
(ADF1, ADF2)—first and second adfrontal setae 
(C1, C2)—clypeal setae 
(D1, D2)—first and second dorsal setae 
(F'1)—frontal seta 
(L1, L2, L3)—first, second, and third lateral setae 
(P1, P2)—first and second posterior setae 
(SD1, SD2)—first and second subdorsal setae 
(SV1, SV2, SV3)—first, second, and third subventral setae 
(XD1, XD2)—first and second prothoracic dorsal setae 


Journal of the Lepidopterists’ Society 
87(3), 1983, 206 


OBITUARY 
JOSEPH MULLER (1903-1982) 


Joseph Muller of Lebanon, New Jersey died in September 1982 at the age of 79 years. 

Joe had pretty much devoted a lifetime to Lepidoptera. He came to this country from 
his native Alsace-Loraine as a young adult and spent the remainder of his life in New 
Jersey. He was always identified with the Newark Entomological Society and indeed for 
years the Fall meeting of that group was held at his home. He loved to recall the society 
beer fests of an earlier era. 

Joe was an authority on the butterflies and moths of New Jersey and frequently 
reported new records, particularly moths, from our state. He authored many articles, 
some of which appeared in the Journal of the Lepidopterists’ Society. In recent years, 
one of his favorite haunts was the farm of Dr. Brooke Worth in Eldora, Cape May 
County, which has been featured in national publications of the Nature Conservancy 
and Audubon Society. 

The Muller collection has been donated to the American Museum of Natural History. 
His spirit and interest in insects and his zest for the chase live on with those who knew 
him well. 


JOHN J. Bowe, Van Neste Medical Arts Center, 127 Union Street, Ridgewood, New 
Jersey 07450. 


Journal of the Lepidopterists’ Society 
37(3), 1983, 207-216 


VARIATION AND HOST SPECIFICITY IN THE YUCCA 
MOTH, TEGETICULA YUCCASELLA (INCURVARIIDAE): 
A MORPHOMETRIC APPROACH 


NANCy JO MILEs’ 


Department of Biology, New Mexico State University, 
Las Cruces, New Mexico 88003 


ABSTRACT. Moths presently recognized as Tegeticula yuccasella Riley were col- 
lected from flowers of Y. baccata, Y. elata, and Y. torreyi from Dona Ana County, New 
Mexico. All three yucca species occurred sympatrically and only Y. elata bloomed later 
than the other two. A morphometric analysis, using characters of the wings and genitalia 
and color of the antennae, was carried out on 225 individuals. The data were separated 
into three groups based on plant host species and analyzed using a stepwise discriminant 
analysis. Significant separation between the three groups resulted, with only three mis- 
classifications. Three presumed separate taxa were described and compared based on the 
analysis, although it was not possible from this study to determine their relationship to 
nominotypic T. yuccasella. For this reason, no new specific names are given. 


Mutualistic relationships between plants and insect pollinators occur 
frequently in nature. Insect dependence on the plant for reasons other 
than a nectar source, however, are much less common. The relationship 
between the orchid Stanhopea and Euglossinae bees, and that between 
wasps of the family Agaonidae and fig trees, are both examples of 
highly specific pollination relationships (Dressler, 1968; Ramirez, 1969). 
In these relationships, specific pollinators exist for closely related species 
of plants and maintain genetic isolation between the species with which 
they are sympatric. These highly specific pollination relationships pro- 
vide a strong basis for speciation and genetic isolation (Baker, 1968). 

The mutualistic relationship between the yucca plant and the yucca 
moth is well known (Riley, 1872; Trelease, 1898; and Rau, 1945). The 
female moth, with its specialized mouthparts for depositing pollen in 
the stigma of the pistil, is the only effective pollinator of yuccas. The 
moth does not obtain nectar from the plant but is dependent upon it 
for the development of its larvae. The female deposits eggs in the ovary 
of a flower where the larvae feed on seeds until reaching maturity. 

In contrast to the examples above, specific yucca moth pollinators 
are not known for sympatric and closely related yucca plants, except 
in yucca moths which pollinate Californian yuccas. Only two moth 
species are thought to pollinate all other species of yuccas in the United 
States and Mexico. One of these, Parategeticula pollenifera Davis, is 
known only in southeastern Arizona and Mexico (Davis, 1967). The 
other moth, Tegeticula yuccasella Riley, as presently recognized, is 
widely distributed across the United States and Mexico. 


1 Present address: Statistical Computing Support Group, 50 Warren Hall, Cornell University, Ithaca, New York 14853. 


208 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Davis (1967) found that variation in the wings and genitalia of T. 
yuccasella was greater than in any other member of the subfamily 
Prodoxinae. Davis examined individuals from 11 species of yucca across 
the geographic range of T. yuccasella and found that differences in 
the wing measurements could be ascribed to geographical variation 
but that differences in the genitalia could not. The variation in the 
genitalia showed a high correlation with plant host species. Davis pro- 
posed that possibly a specific pollinator had evolved for each of the 
four sections (subgenera) of Yucca. Yucca brevifolia Engelm. and Y. 
whipplei Torr., which belong to the sections Clistocarpa and Hespe- 
royucca, respectively, are found in California and have specific polli- 
nators. Davis postulated that T. yuccasella was a specific pollinator for 
the section Chaenocarpa and that possibly a sibling species is in the 
process of evolving for the section Sarcocarpa. 

Southern New Mexico presents an excellent opportunity to examine 
variation in T. yuccasella due to host specificity for two reasons. First, 
three species of yucca representing two sections of the genus occur 
sympatrically in the area. Yucca baccata Torr. and Y. torreyi Shafer 
are members of the section Sarcocarpa and Y. elata Engelm. is a mem- 
ber of the section Chaenocarpa. Second, this sympatry minimizes the 
occluding effects of geographical variation, and one can concentrate 
on variation due principally to plant host specificity. 

Basic differences that exist in the ecology and morphology of these 
three yucca species may be important when considering host specificity 
in plant pollination. The flowers are different in position on the plant, 
shape, odor, and time of flowering. Y. elata bloomed approximately 
one month later than Y. baccata and Y. torreyi, which were blooming 
simultaneously. Since adult yucca moths emerge when yuccas are first 
blooming, it was possible to collect adults from Y. baccata and Y. 
torreyi at the same place and time; whereas, adults from Y. elata were 
collected at the same place but at a different time. 

Two possibilities are being explored: either 1) a specific pollinator 
has developed for the section Chaenocarpa and another for the section 
Sarcocarpa; or 2) a specific pollinator exists for each of the three species 
of yucca in southern New Mexico. The latter implies that more than 
one moth species pollinates the section Sarcocarpa. 


MATERIALS AND METHODS 


T. yuccasella were collected from newly opened flowers of yuccas 
from seven cities in the Las Cruces area of Dona Ana County, New 
Mexico, during the spring of 1979. All collecting was done during the 
day when individuals were inside flowers and could be easily captured. 
Moths were normally only found in newly opened flowers; therefore, 


VOLUME 87, NUMBER 3 209 


VINICULUM 
WIDTH 


LENGTH 


SACCUS 
WIDTH 


LENGTH 


Ss see any 


SIGNUM 
DIAMETER 


Fic. 1. Morphometric measurements taken from T. yuccasella: A, right forewing; 
B, male genitalia; C, aedeagus; D, female genitalia. 


the number of suitable flowers at a given site on a given day was 
limited. An attempt was made to remove every moth from each plant 
found to be harboring moths. Y. baccata was first noted in bloom on 
10 April, and had finished blooming by the last week in April. The 
first Y. torreyi seen in bloom was 14 April, and the last finished bloom- 
ing by the end of April. Y. elata did not begin blooming until the third 
week in May although a few were observed in bloom until the end of 
June. 

All specimens used in the morphometric analysis are deposited in 
the Museum of Entomology, Department of Biology, New Mexico State 


210 


TABLE l. 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Tegeticula collected from Y. baccata, Y. torreyi, and Y. elata. 


Means and standard deviations (mm), with sample sizes in parentheses, for 


Males Y. baccata Y. torreyi Y. elata 
Wing length 10.64 + 0.46 (79) 11.43 + 0.49 (67) 10.08 + 0.63 (89) 
Wing width 2.70 + 0.20 (79) 3.05 + 0.18 (67) 2.85 + 0.22 (89) 
Length of genitalia 2.50 + 0.09 (52) 1.92 + 0.09 (48) 1.53) = Oda G49) 
Width of vinculum 0.80 + 0.08 (87) 0.90 + 0.09 (30) 0.88 + 0.12 (34) 
Width of saccus 0.15 + 0.08 (41) 0.20 + 0.08 (46) 0.18 + 0.04 (48) 
Length of aedeagus 2.85 + 0.14 (51) 2.19 + 0.12 (48) 1:72 = OAS 6O) 

Females 
Wing length 12:73 = 0161 (75) 13.28 = 01545(66) ae aor==. Oa anoo) 
Wing width SJL Se OAL (7s) 3.47 + 0.18 (66) 3.21 + 0.20 (86) 
Diameter of signum 0.40 + 0.08 (24) 0.82 + 0.10 (24) 1.04 + 0.11 (28) 
Height of ovipositor 0.10 + 0.02 (17) 0.138 + 0.02 (19) 0.09 + 0.02 (21) 


University, Las Cruces. Genitalia were removed from 150 males and 
75 females and prepared following the method described by Burns 
(1964). 

Characters were measured using a dissecting microscope with an 
ocular micrometer. Six quantitative characters for males and four for 
females were measured (Fig. 1). Male characters included the maxi- 
mum length of the right forewing, width of the right forewing (as 
measured from a point perpendicular to the point where the cubitus 
of the hind wing branches), genitalic length (defined as the maximum 
length of the vinculum, to the tip of the saccus), width of the saccus, 
width of the vinculum, and length of the aedeagus. Female characters 
included the same two wing measurements plus the diameter of the 
signum including rays and height of the base of the ovipositor. The © 
genital nomenclature follows Klots (1956). In addition to these quan- 
titative characters, a single qualitative character was analyzed: in both 
males and females the color of the antennae. Seven categories were 
established: two categories were all yellow or all brown and the re- 
maining five were bicolored as follows: white-black, light brown-black, 
white-yellow, white-brown, and light brown-yellow. 

Six groups were obtained by separating according to the three host 
species and sexes of the insects. All analyses and programs used were 
of the BMDP series developed by the Health Science Computing Fa- 
cility, University of California (Dixon, 1975). Basic statistics (BMDP1D) 
were carried out for each group to obtain means and standard devia- 
tions. Each of the six groups was subdivided by collecting sites and 
analyzed via analysis of variance (BMDPI1V) to test for differences in 
means among localities. This was carried out to ensure that geograph- 


VOLUME 87, NUMBER 3 Dial 


SECOND CANONICAL VARIABLE 


aS 30 OO 30 U9) 


FIRST CANONICAL VARIABLE 


Fic. 2. Plot of the first two canonical variables from the stepwise discriminant anal- 
ysis for 91 Tegeticula males. Numbers represent centroids; E represents individuals 
collected from Y. elata, T represents individuals collected from Y. torreyi, and B rep- 
resents individuals collected from Y. baccata. 


ical variation was nonsignificant (P < 0.05). After these preliminary 
analyses, separate discriminant analyses (BMDP7M) for males and fe- 
males were used for determining discrimination among the three plant 
host groups. The discriminant function used is based on the Maha- 
lanobis D? value, a measure of the metric distance between population 
centroids (Atchley & Bryant, 1975). 


RESULTS 


Two hundred and twenty-five moths were measured (Fig. 1). Wing 
and genitalic characters were quite distinct for each plant host group 
and are given in Table 1. 

Fig. 2 shows the plot of the first two canonical variables for each of 
the three groups of males. The variables entered in their order of 
significance were length of the genitalia, wing length, length of the 
aedeagus, and width of the saccus. Of 91 individuals (the remaining 
cases were dropped because they contained missing data) there were 
only two misclassifications, giving an overall percent correct classifi- 
cation of 98.9. Misclassifications are circled on the figure. 


Dale JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


SECOND CANONICAL VARIABLE 


6.0 3.0 0.0 30) 6.0 


FIRST CANONICAL VARIABLE 


Fic. 3. Plot of the first two canonical variables from the stepwise discriminant anal- 
ysis for 54 Tegeticula females. Numbers represent centroids; E represents individuals 
collected from Y. elata, T represents individuals collected from Y. torreyi, and B rep- 
resents individuals collected from Y. baccata. 


Fig. 3 shows the plot of the first two canonical variables for each of 
the three female groups. Signum diameter, antennal color, ovipositor 
height, and wing length were the characters entered in their order of 
significance. There was only one misclassification out of 54 individuals, 
giving an overall correct classification of 98.1%. 

Table 2 shows the eigenvalues and coefficients for the first two ca- 
nonical variables for both sexes. The first canonical variable is the linear 
combination of characters added that best discriminates among the 
groups. 

A separation of groups of this magnitude with such little misclassi- 
fication is strong evidence that three distinct taxa are present, especially 
when coupled with the host-specificity and sympatry data. It appears 
that three taxa of moths exist in southern New Mexico among popu- 
lations previously considered to represent one species, T. yuccaselia. 
One of the taxa revealed by the discriminant analyses may represent 
a geographical component of nominotypic T. yuccasella. Further re- 


VOLUME 37, NUMBER 3 2S 


TABLE 2. Eigenvalues and coefficients for the first two canonical variables for males 
and females based on stepwise discriminant analysis. 


Females 
Coefficients for canonical variables 
Variable ] 2 
Wing length —0.00364 — 0.12587 
Antennae color — 1.23943 0.69803 
Signum diameter — 0.27162 —0.15915 
Ovipositor height 0.37599 — 0.88764 
Constant term 8.37599 16.0337 
Eigenvalue 12.0282 1.55944 
Males 
Coefficients for canonical variables 
Variable 1 2 
Wing length —0.04152 —0.21477 
Genitalia length 0.23821 0.12548 
Aedeagus length 0.11516 —0.09967 
Genitalia width —0.19504 —0.49490 
Constant term — 17.9077 17.9449 
Eigenvalue 21.6712 0.89412 


search needs to be conducted to determine the hierarchic ranking of 
the three taxa. Therefore, I include a diagnosis of each below without 
proposing formal names. 


COMPARISON OF TAXA STUDIED 


Tegeticula ex Y. baccata: Head white, with antennae usually white for the first half of 
length and black to tip, or occasionally antennae appear completely black due to wear; 
maxillary tentacle fully developed and labial palpus brown; thorax white; forewings 
white dorsally except for almost black fine line along proximal half of length of costal 
vein; forewings dark gray ventrally except for white fringe; length of the forewings 9.43 
to 11.7 mm in males and 11.43 to 14.14 mm in females; hind wings dark gray dorsally, 
hind wings dark gray ventrally for costal one-third of length and lighter posterior two- 
thirds; fringe of hind wing gray, occasionally white; abdomen pale brown dorsally and 
white ventrally. Genitalia (Fig. 4): tegumen bilobed with lobes widely separated; saccus 
elongate and narrow; aedeagus elongate and slender, length 2.4 to 3.1 mm; ovipositor 
with convex minutely serrate ridge with slightly more than 30 teeth; height of the base 
of the ovipositor intermediate between other two species; signum and rays small with 
diameter 0.32 to 0.47 mm; number of rays exceeds 20. 


This form was only collected from newly opened flowers of Y. bac- 
cata with the exception of six individuals collected on Y. torreyi flow- 
ers. These six individuals emerged after all available Y. baccata flowers 
were gone and Y. torreyi was the only yucca in bloom. 

Tegeticula ex Y. elata: Head white, with antennae brown, yellow or bicolored white- 


brown; maxillary tentacle usually fully developed and labial palps brown; thorax white; 
forewings white dorsally except for fine black line along costal vein; forewings tan ven- 


214 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Fic. 4. Drawings of male genitalia: A, Tegeticula ex Y. torreyi; B, Tegeticula ex Y. 
elata; C, Tegeticula ex Y. baccata. 


trally except for white fringe; occasionally white extends into margin of wing slightly; 
length of forewings 8.57 to 11.86 mm in males and 10.07 to 18.29 mm in females; hind 
wings tan to almost white dorsally; hind wings brown ventral costal one-third of length 
and white posterior two-thirds; fringe on hind wings white; abdomen white dorsally and 
ventrally. Genitalia (Fig. 4): tegumen bilobed with lobes widely separated; saccus much 
shorter and wider than in Tegeticula ex Y. baccata; aedeagus also much shorter, length 
1.4 to 2.0 mm; ovipositor with minutely serrate ridge with slightly more than 30 teeth; 
height of the base of the ovipositor lowest of the three species; signum and rays largest 
in diameter of the three species, diameter 0.90 to 1.30 mm; number of rays exceeds 20. 


This form was collected in newly opened flowers of Y. elata. This 
was the only available yucca species in bloom at the time these moths 
were active. 


Tegeticula ex Y. torreyi: Head white, with antennae yellow or bicolored white-yellow; 
maxillary tentacle fully developed and labial palpus brown; thorax white; forewings 
white dorsally except for fine black line along costal vein; forewings gray ventrally with 
white fringe; gray lighter than in Tegeticula ex Y. baccata; length of forewings 10.48 to 
13.00 mm in males and 12.14 to 14.57 mm in females; hind wings light gray ventrally, 
costal one-third of wing darker gray than posterior two-thirds; abdomen pale brown 
dorsally and white ventrally. Genitalia (Fig. 4): tegumen bilobed with lobes widely 
separated; saccus elongate but not as much as in Tegeticula ex Y. baccata; saccus wider 
than Tegeticula ex Y. baccata; aedeagus elongate and slender, length 2.0 to 2.8 mm; 
ovipositor with minutely serrate ridge with slightly more than 30 teeth; height of the 
base of the ovipositor intermediate between the other two species; signum and rays only 
slightly more narrow than Tegeticula ex Y. elata, diameter 0.43 to 0.93 mm; number of 
rays exceeds 20. 


This form was collected from newly opened flowers of Y. torreyji. Y. 


VOLUME 387, NUMBER 3 215 


baccata was also in bloom at the same time but no individuals were 
collected from that plant. 


Species Diagnosis 


The characters that are best for distinguishing among the three moths 
are antennal color, wing color, abdomen color, male genitalia length 
and width, aedeagal length, signum diameter, and height of the base 
of the ovipositor. The genitalia and wing measurements given by Davis 
(1967) for T. yuccasella are highly variable and overlap those of these 
three moths in each case and cannot be used to separate them. It should 
be noted that the measurements taken by Davis included individuals 
collected from Y. baccata, Y. elata, and Y. torreyi in southern New 
Mexico and probably do not all represent T. yuccasella. 


DISCUSSION 


The results of this study support the hypothesis that a single species 
of moth exists for each of the three species of yucca in southern New 
Mexico. The data do not support Davis’ hypothesis (1967) that one 
species exists for each of the sections of the genus Yucca. Tegeticula 
ex Y. baccata and Tegeticula ex Y. torreyi both have hosts within the 
section Sarcocarpa and are quite distinct morphologically. T. yucca- 
sella as described by others, then, is a composite species. The study 
also implicates that in the Las Cruces area, one of these three moths 
may represent a subspecies of nominotypic T. yuccasella. To answer 
this question, additional studies similar to this one need to be carried 
out at other localities between southern New Mexico and Missouri (the 
collection locality for the holotype of T. yuccasella). Moreover, anal- 
yses of yucca moths and their hosts over broader geographic ranges 
may identify additional species of Tegeticula. 

Justification for assigning specific status to these three taxa would 
stem from morphological distinctness and fidelity to species of yucca. 
Samples collected from the same locality can be either individual vari- 
ants of the same species or else different species (Mayr, 1969). In de- 
ciding between the two alternatives, it is important to examine char- 
acters such as the genitalia which are not subject to a great deal of 
individual variation and look for intermediacy between the groups. If 
differences among groups are found consistently in unrelated charac- 
ters, specific rank is further implicated (Mayr, 1969). All three groups 
from this study are sympatric and show differences in the genitalia, as 
well as other very different characters such as wing length and antennal 
color. No individuals with intermediate characters were found and this 
indicates an absence of gene flow, particularly between moths collected 


216 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


on Y. baccata and Y. torreyi, where blooming times were simultaneous. 
Temporal isolation appears to exist among the Y. elata group and the 
two previous groups, because blooming times and thus time of moth 
emergence are a full month apart. Since moths only live two to three 
days, overlap between breeding periods is not possible. Therefore, 
blooming times serve as a principal mechanism for isolating the Y. 
elata group from the other two groups in the area. Following the above 
criteria, moths taken from the three plant host species are themselves 
distinct species. 

Six Tegeticuia ex Y. baccata were collected from flowers of Y. tor- 
reyi. Two of these were included in the discriminant analysis and 
account for misclassifications. These moths emerged in an area where 
all the Y. baccata had finished blooming and Y. torreyi was the only 
available flowering yucca. This shows that, although moths usually 
maintain host specificity, individuals potentially can and do use alter- 
nate yucca hosts if the preferred host is unavailable. The fact that these 
individuals were observed in the presence of Tegeticula ex Y. torreyi 
further supports the conclusion that there is no gene flow between the 
two because no intermediates were observed. 


ACKNOWLEDGMENTS 


I want to thank Dr. James R. Zimmerman for his continual support and guidance and 
Gregory S. Forbes for his field assistance. 


LITERATURE CITED 


ATCHLEY, W. R. & E. H. BRYANT. 1975. Multivariate Statistical Methods: Among- 
groups Covariation (Volume 1). Douden, Hutchinson and Ross, Inc., New York. 
BAKER, H. G. 1963. Evolutionary mechanisms in pollination biology. Science 139:877— 
883. 

BuRNS, J. M. 1964. Evolution in skipper butterflies of the genus Erynnis. Univ. Cali- 
fornia Pub. Entomol. 37. iv + 230 pp. 

Davis, D. R. 1967. A revision of the moths of the subfamily Prodoxinae (Lepidoptera: 
Incurvariidae). United States Nat’] Mus. Bull. 255. vii + 170 pp. 

Dixon, W. J. 1975. Biomedical Computer Programs. Univ. California Press, Berkeley 
and Los Angeles. 

DRESSLER, R. L. 1968. Pollination of Euglossinae bees. Evolution 22:202-210. 

Kuots, A. B. 1956. Lepidoptera. In S. L. Tuxen (ed.). Taxonomic Glossary of Genitalia 
of Insects. S. H. Service Agency Inc., Darien, Connecticut. 

Mayr, E. 1969. Principles of Systematic Zoology. McGraw-Hill Book Company, New 
York. 

RAMIREZ, W. 1969. Fig wasps: Mechanism of pollen transfer. Science 163:580-581. 

Rau, P. 1945. The yucca plant, Yucca filamentosa, and the yucca moth, Tegeticula 
(Pronuba) yuccasella Riley: An ecologico-behavior study. Ann. Missouri Bot. Garden 
32:373-394. 

RILEY, C. V. 1872. The fertilization of the yucca plant by Pronuba yuccasella. Can. 
Entomol. 4:182. 

TRELEASE, W. 1893. Further studies of yuccas and their pollination. Fourth Ann. Rep. 
Missouri Bot. Garden 4:181-226. 


Journal of the Lepidopterists’ Society 
37(3), 1983, 217-223 


IMMATURE STAGES OF ANACAMPTODES HERSE (SCHAUS) 
(GEOMETRIDAE) ON SOYBEAN IN HONDURAS 


STEVEN PASSOA 


Department of Entomology and Nematology, University of Florida, 
Gainesville, Florida 32611 


ABSTRACT. Anacamptodes herse is recorded from Honduras for the first time. The 
larvae were reared on soybean and are described with the pupa. 


The genus Anacamptodes contains 24 species distributed throughout 
the New World from Canada to Costa Rica (Rindge, 1966; McGuffin, 
1977). Three species of Anacamptodes are recorded as pests (Dixon, 
1982; Furniss & Carolin, 1977; Zimmerman, 1958), but none is a prob- 
lem in Honduras. Geometrids are important defoliators of soybeans in 
other parts of the world. Panizzi et al. (1980) listed four genera at- 
tacking soybeans in Brazil (Iridopsis, Oxydia, Semiothisa, and Sten- 
alcidia). Park et al. (1978) considered six genera important in Japan 
(Ascotis, Biston, Bizia, Scopula, Serraca, and Ectropis). 

Rindge (1966) reported Anacamptodes herse (Schaus) from Mexico 
and Costa Rica while stating the immature stages were unknown. This 
paper describes the mature larva and pupa of A. herse and establishes 
its occurrence in Honduras. Larvae were initially swept from and reared 
on soybean, Glycine max (L.) Merrill, outside the city of La Paz, de- 
partment of La Paz, Honduras. These larvae were associated with a 
high population of Anticarsia gemmatalis Hbn. (Noctuidae) but were 
not causing economic damage. 


METHODS AND MATERIALS 


The larvae were described from three shed skin preparations with 
associated adults in addition to larvae preserved in alcohol. A male 
(genitalia slide #159, shed skin and mandible slide #166, S. Passoa 
coll.) and two females (genitalia slides #145, 285, shed skin and man- 
dible slides 116, 221, S. Passoa coll.) were examined. Color photographs 
also aided in the descriptions. 

Collection data are as follows: Honduras, Jutiapa (near Danli), 1-IX-1979, larva on 
soybean leaf, coll. E. M. de Vasquez, mandible slide #169, S. Passoa coll.; Honduras, La 
Paz (near Comayagua), 14-VIII-1980, larva on soybean, mandible slide #170, S. Passoa 
coll., male and female emerged 28-VIII-1980, coll. S. Passoa; Honduras, E] Zamorano, 


Escuela Agricola Panamericana, 9-IX-1982, larva on soybean, female emerged 24-IX- 
1982, coll. S. Passoa. 


The shed skins of the last instar larvae after pupation were first — 
softened in 10% potassium hydroxide for 24 hours. Later they were 
washed in acid alcohol followed by increasing alcohol concentrations 


218 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


until absolute alcohol was reached. At this point the head was removed. 
The epicrania, frontal area, mandibles, and hypopharyngeal complex 
were separated from each other. Slide mounted material was cleared 
in clove oil and mounted in balsam. Two of the three shed skins were 
stained in mercurochrome and slide mounted. The other was preserved 
in alcohol. 


RESULTS 


Larva: Coloration of living material: head reticulated with light brown spots on a 
cream ground, top of vertex tipped with brown, middle of frontal area often with a 
black spot, body faintly lined with long dark green-brown and white stripes on a light 
green ground, thoracic legs and all spiracles both tan-brown, A2 with a black dorsal 
projection usually tipping seta D2, often another black spotted tubercle on A2 behind 
and below the spiracle touching L1, D1 of A8 tuberculate and black (Figs. 1, 12). 

Early instar larva: yellow-green with a tan head, dorsal tubercle of A2 present but 
reduced in size, lateral tubercles absent. 

Chaetotaxy: Head with P2 directly above P1, A2 in front and below A8 and above 
Al, L1 above O2, O1 centrally located between stemmata, adfrontals with AF] and AF2 
widely separated, Fl above and behind C2, Cl close to C2, labrum with L1 and L2 
longer than L38, M1, M2, and M8 as shown (Figs. 7, 8). Prothorax with XD1, XD2, D1, 
and D2 approximately equidistant from each other, SD1 and SD2 closely spaced, Ll 
above and in front of L2, SV group bisetose (Fig. 10). Mesothorax with D1, D2, SD1, 
and SD2 in a vertical line, D2 and SD2 widely separated, L3 longer than L1 or L2, one 
SV seta present (Fig. 11). Second abdominal segment with D1 and D2 level with each 
other, SD1 in front of the spiracle, L1 behind the spiracle, L2 above L8, SV2 absent, 
SV3 above and behind SV4, SV4 above SV1 (Fig. 12). Sixth abdominal segment with D1 
above D2, SD1 in front of spiracle, L1 behind it, L2 directly above L8 (Fig. 18), five SV 
setae present on A6 proleg. Eighth abdominal segment with D setae as in A6, SD1 closer 
to the spiracle, L1 equidistant from L2 and SV3, one SV seta present (Fig. 14). Ninth 
abdominal segment with D1 in front and below D2, SD1 above L1, L1 above SV1 (Fig. 
14). Tenth segment with SD1 and D1 widely spaced, L1 and D2 close together, all four 
setae on the anal shield, CD1 above CD2, both on the paraproct, LG3 above LG2 and 
LG1, CP1 above CP2 (Fig. 14). One ventral seta is present on each segment, SV2 absent 
on Al (Fig. 20). | 

Mouthparts: mandible shape variable, usually with eight teeth (Figs. 2, 3), sometimes 
all worn smooth (Fig. 4), first four teeth larger than the others, lateral tooth and two 
mandibular setae present. Hypopharyngeal complex with labial palps about as long as 
spinneret, small stipular setae present, proximomedial region membranous sparsely cov- 
ered with fine spines (Figs. 5, 9). 

General: last instar larva about 22 mm long, setae arising from small black chalazae, 
skin granulated with short truncate cones (Fig. 18), tarsal claw rounded with three 
clubbed setae at its base (Fig. 6), A6 with crochets incompletely formed into two groups - 
in early instar larvae (Fig. 19). Mature larvae with biordinal mesoseries in unbroken 
band. 

Pupa: With large eyes, semicircular labrum, oblong labial palps, and maxillae extend- 
ing with antennae to caudal margin of wings, prothoracic femur exposed, prothoracic 
leg extending about % length of maxillae, mesothoracic leg ending near antennae (Fig. 
15). Prothoracic callosity oval and spinose (Fig. 16). Cremaster with two diverging spines 
forming “V” between them, usually broken (Fig. 17), but each spine actually bifurcate 
at tip. Abdominal segments punctate. Length varies, 12 mm (male) to 16 mm (female), 
colored reddish brown. 


DISCUSSION 


The position of the protuberances on A2 is an important character 
separating Anacamptodes from its close relatives (Heitzman, 1982), 


_ VOLUME 387, NUMBER 3 219 


Fics. 1-6. Anacamptodes herse larva: 1, dorsolateral view of mature larva showing 
defoliation of soybean leaf in the background; 2, mandible with all teeth sharp; 3, 
mandible with teeth partially worn; 4, mandible with teeth worn smooth; 5, hypopha- 
ryngeal complex; 6, tarsal claw and its setae. (Scale line = 5.5 mm, 0.4 mm, 0.38 mm, 0.3 
mm, 0.4 mm, 0.5 mm, respectively) 


220 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Fics. 7-9. Anacamptodes herse larval head structures: 7, epicrania; 8, adfrontal area 
and labrum; 9, hypopharyngeal complex; scale line = 0.25 mm, 0.25 mm, and 0.15 mm, 
respectively. (A = anterior setae; AF = adfrontal setae; C = clypeal setae; F = frontal se- 
tae; L = lateral setae; LP = labial palps; M = medial setae; O = ocular setae; P = posterior 
setae; S = spinneret) 


but their size and shape is variable and may be missing in early instars. 
Younger larvae of A. herse have the D2 protuberance reduced in size 
and the L2 protuberance absent. Similar results were reported by Com- 
stock and Dammers (1946) studying Anacamptodes fragilaria (Gross- 


— 


Fics. 13, 14. Anacamptodes herse chaetotaxy continued: 13, Anterior portion of A6; 
14, Segments 8, 9, and 10. Scale line = 0.38 mm. (CD = dorsal coxal setae; CP = posterior 
coxal setae; D = dorsal setae; HYP = hypoproct; L = lateral setae; LG = lateral setae of 
the proleg; PRP = paraprocts; SD = subdorsal setae; SP = spiracle; SV = subventral setae) 


VOLUME 37, NUMBER 3 22 


10 11 42 


——4 


Fics. 10-12. Anacamptodes herse chaetotaxy: 10, prothorax; 11, mesothorax; 12, 
2nd abdominal segment; scale line = 0.3 mm. (A = abdominal segment; D = dorsal setae; 
L = lateral setae; SD = subdorsal setae; SP = spiracle; SV = subventral; XD = prothoracic 
dorsal setae) 


222 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Fics. 15-20. Details of Anacamptodes herse larva and pupa: 15, pupa, ventral view; 
16, prothoracic callosity; 17, pupal cremaster; 18, skin of larva at 400x; 19, crochets 
of A6; 20, ventral view of Al and AQ; scale line = 1.4 mm, 0.4 mm, 0.7 mm, 0.025 mm, 
0.4 mm, and 0.5 mm, respectively. (SV = subventral setae; V = ventral setae) 


beck). Only the last instars have the protuberances present, earlier 
instars lack them. Two separate rearings of the related A. defectaria 
(Gn.) from Gainesville, Florida, show a variation in the size and shape 
of D1 on A2. Some have a large upright conical protuberance, while 
others show a more flattened ridge-like structure. Given the above 
variation in the genus, it would be difficult to generalize and present 
the typical form of A. herse without a large series of larvae. McGuffin 
(1967) described Anacamptodes angulata Rindge from Mexico on Ar- 
temisia sp. as having “a dorsal ridge on A2 between setae D2, seta D1 
on A8 prominently tuberculate,” which is similar to the situation in A. 
herse. Color pattern can also be variable. Furniss and Barr (1967), 
cited by McGuffin (1977), reported four color forms of A. clavinaria 
profanata (Barnes and McD.) in the field. This variability in structure 
and color makes specific identification difficult. McGuffin (1977) relied 


VOLUME 387, NUMBER 3 DO 


on distribution and hosts to separate the Canadian species of Ana- 
camptodes instead of larval morphology. 

The chaetotaxy is also very variable. The following setae can be 
more anterior than illustrated: Al (head); SV2, SV1 (T1); D1, SV4, SV1 
(Al); and L8 (A6). These setae can be more posterior than illustrated: 
P setae (head); D2 (Al); SD1 (A6) and SV1 (A8). The spiracle of A6 
may lie directly above L2, more posterior than shown. M2 and L3 (T2) 
can be more dorsad, L1 (A6, A8) more ventrad, than the normal po- 
sition. Finally, the AF, SD (T1), and L (T1) setae can be closer to each 
other than illustrated. 

Little information is available for other tropical species of Anacamp- 
todes, but an important similarity is shared by the Canadian species 
of Anacamptodes and A. herse concerning the relative length of the 
D setae on A3 compared to the spiracle size on that segment. In both 
cases the D setae are as long, or longer, than the spiracle. 

It is worth noting that an as yet unidentified Anacamptodes was 
reared on soybean at Zamorano, Honduras. Unfortunately, no infor- 
mation is available on its early stages. Therefore, collectors wishing a 
positive specific determination must depend on rearing an adult male. 
The male genitalia of A. herse are distinctive (Rindge, 1966). 


ACKNOWLEDGMENTS 


I wish to thank E. M. de Vasquez for the larval specimen used in this study. N. Urbina 
and M. Funez provided transportation and advice on where to find soybean insects. 
Thanks are also due to G. Reyes who provided office space at Recursos Naturales in 
Comayagua, Honduras. This study was supported by Peace Corps, Honduras. Dr. Rindge 
determined the adult male. The comments of D. Habeck and R. Heitzman on the 
manuscript were most helpful. Florida Agr. Journal Series no. 4851. 


LITERATURE CITED 


COMSTOCK, J. A. & C. M. DAMMERS. 1946. Notes on the life history of Anacamptodes 
(Cleora) fragilaria (Grossb.). Bull. So. Calif. Acad. Sci. 45(1):17-20. 

Dixon, W. N. 1982. Anacamptodes pergracilis (Hulst), a cypress looper. Fla. Dept. 
Agr. & Consumer Serv. Entomol. Cir. no. 244. 2 pp. 

FURNISS, R. L. & V. M. CAROLIN. 1977. Western forest insects. USDA For. Serv. Misc. 
Pub. 275. 

HEITZMAN, R. L. 1982. Descriptions of the mature larva and pupa of Hypomecis 
umbrosaria (Hbn.). Proc. Entomol. Soc. Wash. 84(1):111-116. 

McGurFFIn, W. C. 1967. Immature stages of some Lepidoptera of Durango, Mexico. 
Can. Entomol. 99:1215-1229. 

1977. Guide to the Geometridae of Canada II. Subfamily Ennominae 2. Mem. 
Entomol. Soc. Can. 101. 191 pp. 

PANIzzI, A. R. & B. S. CORREA. 1980. Geometrideos em soja: flutuacao estacional e 
ressurgencia apos o uso de insecticidas. Pesq. Agropec. Brasil 15(2):159-161. 

PaRK, K. T., C. Y¥. HWANG & K. M. CHol. 1978. Lepidopterous insect pests on soybeans. 
Kor. J. Pl. Prot. 17(1):1-5. 

RINDGE, F. H. 1966. Revision of the moth genus Anacamptodes. Bull. Amer. Mus. 
Nat. His. 132:175-244. 

ZIMMERMAN, E. C. 1958. Insects of Hawaii-Macrolepidoptera. Univ. of Hawaii Press, 
Honolulu. Vol. 7, 542 pp. 


Journal of the Lepidopterists’ Society 
87(3), 1983, 224-227 


NEW HOST RECORDS FOR OLETHREUTINAE 
(TORTRICIDAE) 


RICHARD L. BROWN,’ J. F. GATES CLARKE,” AND DALE H. HABECK® 


ABSTRACT. Host records are given for 33 species of Nearctic Olethreutinae in the 
genera Epiblema, Eucosma, Pelochrista, Phaneta, Cydia, Ecdytolopha, Ethelgoda, 
Grapholita, Satronia, Episimus, Larisa, and Zomaria. 


Host plant records have been published for nearly half of the ap- 
proximately 800 species of Nearctic Olethreutinae. The following rec- 
ords are given for 33 species represented in the U.S. National Museum 
of Natural History and the Florida State Collection of Arthropods; 
specimens in the latter are indicated by FSCA following the entries. 
Identifications of the Olethreutinae were made by the senior author. 
Plant identifications have not been confirmed since their original iden- 
tifications; host information is given as recorded on the label, except 
the nomenclature has been emended following Kartesz and Kartesz 
(1980). Family names of hosts are given after the initial listing of the 
plant genus. The letter (n) represents the number of specimens reared. 
Dates are given as on the specimen labels and do not imply natural 
emergence times because of the various rearing conditions. 

The publication of host records should be tempered with a precau- 
tionary note. The following records do not imply that the listed plant 
species is the favored host. Some species may be incidental hosts, while 
others may represent the plant upon which the larva was collected and 
not necessarily the plant on which the larvae were feeding or the plant 
upon which the female oviposited. More than one larval collection 
record from a particular plant provides greater evidence of the pre- 
ferred host. As some plant species might possibly be misidentified, it 
is here recommended that voucher specimens of host plants be main- 
tained in a reference collection accompanying the reared insect. 


EUCOSMINI 


Epiblema benignata McDunnough 

Artemisia dracunculus L. (Asteraceae)— Washington: Whitman Co., Snake River opp. 
Clarkston, J. F. G. Clarke, gall maker on stem (6n). 

Artemisia vulgaris L.—same data as A. dracunculus (49n). An additional series of 15 
specimens has been reared from A. vulgaris at Wilma, Whitman Co., by Clarke. 
However, these individuals did not form galls on the stems. The reared adults are 
slightly smaller than those from Clarkston, but do not differ in maculation or geni- 
talia. 


1 Department of Entomology, Drawer EM, Mississippi State, Mississippi 39762. 
2 Department of Entomology, Smithsonian Institution, Washington, DC 20560. 
8 Department of Entomology and Nematology, University of Florida, Gainesville, Florida 32611. 


VOLUME 87, NUMBER 3 225 


Epiblema sosana (Kearfott) 

Ambrosia acanthicarpa Hook. (Asteraceae)—CALIFORNIA: Los Angeles Co., Sierra 
Madre (ln); Riverside Co., Sunnymead (6n); San Bernardino Co., Cucamonga (5n); 
R. D. Goeden and D. W. Ricker; AA Lots 69-2A, 69-47A, 69-6B, 69-7B, 68-2B. 

Epiblema scudderiana (Clemens) 

Heterotheca subaxillaris (Lam.) Britt. & Rusby (Asteraceae)—FLORIDA: Alachua 

Co., Gainesville, 22 Aug. 1980, D. H. Habeck, A-2663, FSCA (8n). 
Eucosma agricolana (Walsingham) 

Artemisia vulgaris L.—-WASHINGTON: Whitman Co., em. 8 May-5 Jun. 1932, J. F. 

G. Clarke (10n). 
Eucosma bilineana Kearfott 

Helianthus tuberosus L. (Asteraceae)—ILLINOIS: Du Page Co., Hinsdale, em. 24 

Jan.—24 Feb., Satterthwait (10n). 
Eucosma bolanderana (Walsingham) 

Machaeranthera canescens (Pursh) Gray (Asteraceae)—ARIZONA: Pinal Co., Supe- 

rior, em. 24 Sept. 1937; boring in roots (1n). 
Eucosma mandana Kearfott 

Solidago sp. (Asteraceae)—WASHINGTON: Whitman Co., Almota, em. 15 Mar.-16 

May, J. F. G. Clarke; boring in roots (9n). 
Eucosma ridingsana (Robinson) 

Heterotheca villosa (Pursh) Shinners (Asteraceae)—WASHINGTON: Whitman Co., 
Snake River opp. Clarkston, em. 13 Sept., J. F. G. Clarke (1n). 

“Greasewood’ —COLORADO: El Paso Co., Garden of the Gods; root borer (1n). 

Eucosma totana Kearfott 

Chrysothamnus nauseosus (Pall.) Britt. (Asteraceae)—IDAHO: Lincoln Co., 4 mi. NE 

Richfield, 21 Jul. 1961, W. F. Barr (1n). 
Pelochrista rorana (Kearfott) 

Helianthus annuus L. (Asteraceae)—-WASHINGTON: Whitman Co., Almota, em. 

Jan.—May, J. F. G. Clarke, in roots (12n). 
Phaneta amphorana (Walsingham) 

Grindelia sp. (Asteraceae) WASHINGTON: Whatcom Co., Bellingham, em. 19 Oct.- 

20 Dec. 1932, 1933, J. F. G. Clarke (15n). 
Phaneta argenticostana (Walsingham) 

Artemisia dracunculus L.-WASHINGTON: Whitman Co., Snake River opp. Clark- 
ston, em. 138 May 1982, J. F. G. Clarke (4n); Wilma, em. 22 Mar.-5 May 1935, J. F. 
G. Clarke (8n). 

Phaneta dorsiatomana (Kearfott) 

Artemisia vulgaris L.—WASHINGTON: Whitman Co., Snake River opp. Clarkston, 
em. 31 Jan.-13 Mar. 1933, J. F. G. Clarke (7n); Wilma, em. 18 Feb.-17 Mar. 1934, 
J. F. G. Clarke (18n). 

Phaneta misturana (Heinrich) 

Artemisia tridentata Nutt.—IDAHO: Owyhee Co., 7 mi. S Bruneau, 10 Apr. 1963, O. 
O. Fillmore, 020-19 (1n); Twin Falls Co., 6 mi. W Rogerson, 11 Apr. 1963, W. F. 
Barr, 843-01 (3n). 

Atriplex confertifolia (Torr. & Frem.) S. Wats (Chenopodiaceae)—IDAHO: Cassia 
Co., 6 mi. NE Malta, 22 May 1962, W. F. Barr, 764-01 (1n). 

Phaneta stramineana (Walsingham) 

Haplopappus sp. (Asteraceae)—TEXAS: Cameron Co., Brownsville, May 1945, Lot 45- 
18040, Brownsville 59044 (8n). 

Haplopappus suffructicosus (Nutt.) Gray—ARIZONA: Pima Co., 35 mi. S Tucson, em. 
Oct. 22, 1987, A. Voth and L. P. Wehrle (2n). 


GRAPHOLITINI 


Cydia populana (Busck) 
Populus tremuloides Michx. (Salicaceae)—ARIZONA: Kaibab National Forest. 4 Jun. 
1965, D. A. Pierce, Hopk. #51106-6 (2n). 


226 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Cydia flavicollis Walsingham 

Pithecellobium unguis-cati (L.) Benth. (Fabaceae)—FLORIDA: Sugarloaf Key, 22 
Apr. 1945, Lot 45-8793 (1n). 

P. arboreum (L.) Urb.—PUERTO RICO: Mayaguez, in seeds (2n). 

Cydia garacana (Kearfott) 
Populus sp.—ILLINOIS: Cook Co., Chicago, 16 Jun. 1920, E. Beer (1n). 
Cydia ingrata (Heinrich) 
Fraxinus pennsylvanica Marsh. (Oleaceae)—NORTH DAKOTA: Bottineau Co., 9 Jun. 
1970, M. E. McKnight, Hopk. #54051 (1n). 
Cydia lautiuscula (Heinrich) 
Salix sp. (Salicaceae) WASHINGTON: King Co., Seattle, 25-27 Apr. (6n). 
Ecdytolopha mana (Kearfott) 

Celtis sp. (Ulmaceae)—-TEXAS: Cameron Co., Brownsville, 20 Apr. 1945, Lot 45-9929, 
in leaf gall (1n); Travis Co., Austin, 29 Jun., 3 Aug. 1953, J. Riemann, in petiole gall 
(6n). 

Ethelgoda texanana (Walsingham) 

Stillingia sylvatica Garden ex L. (Euphorbiaceae)—-FLORIDA: Charlotte Co., 18 Sept. 
1949 (In). 

Stillingia sp.—FLORIDA: Polk Co., Lake Alfred, 1 Jun.-16 Jul., L. J. Bottimer (Sn). 

Euphorbia sp.—TEXAS: “Monard,”’ 18-20 Jun., L. J. Bottimer, in seed capsules (2n). 

Grapholita conversana (Walsingham) 

Trifolium douglasii House (Fabaceae)—IDAHO: Idaho Co., Grangeville, em. 6 Apr., 

T. R. Chamberlin and L. P. Rockwood (Qn). 
Grapholita lana (Kearfott) 

Lupinus sp. (Fabaceae)—CALIFORNIA: San Diego Co., Cuyamaca Lake, 27 Apr. 
1935, C. Dammers, on leaves (1n). 

Sophora leachiana M. Peck (Fabaceae)—OREGON: Josephine Co., 6 May 1977, Cher- 
yl Crowder (6n). 

Grapholita lunatana (Walsingham) 
Lathyrus sp. (Fabaceae)—OREGON: Washington Co., Forest Grove, July 1924, S. K. 
Keen (ln); Multnomah Co., Portland, em. 24 Apr.—2 May, S. K. Zimmerman (2n). 
Grapholita packardi (Zeller) 
Pyracantha sp. (Rosaceae)—TEXAS: Jefferson Co., Beaumont, 8-15 Jan. (2n). 
Grapholita vitrana (Walsingham) 

Astragalus sp. (Fabaceae)—CALIFORNIA: San Diego Co., Otay, 12 Mar. 1948, E. D. 
Algert (5n). 

Satronia tantilla Heinrich 

Pinus elliottii Engelm. (Pinaceae)—FLORIDA: Baker Co., em. 16 Apr., Hopk. #40169, 
B. H. Ebel, on male flowers (8n). 

Pinus sp.—ARKANSAS: Johnson Co., “RP,” em. 4 Apr. 1967, I. Brown on male flowers 
(2n). 


OLETHREUTINI 


Episimus tyrius Heinrich 
Prunus caroliniana (Mill.) Ait. (Rosaceae)—FLORIDA: Alachua Co., Gainesville, 21 
May 1973, em. 11 Jun. 1973, D. H. Habeck (In); 30 Jul. 1980, em. 16 Aug. 1980 
(In), FSCA. 
Larisa subsolana Miller 
Carya illinoensis (Wangenh.) C. Koch (Juglandaceae)—TEXAS: Brown Co., Browns- 
wood, 26 May 1919, A. T. Fabis, Quaintance No. 18829 (ln); Menard Co., Ft. 
McKavett, em. 26 Jun. 1919, A. T. Fabis, Quaintance No. 188828 (In). 
Zomaria andromedana (Barnes and McDunnough) 
Lyonia ferruginea (Walt.) Nutt. (Ericaceae)—-FLORIDA: Marion Co., 11 mi. E Lynne, 
17 Sept. 1980, em. 29 Sept.-13 Oct. 1980, D. H. Habeck, A-2684f, FSCA. 
Zomaria interruptolineana (Fernald) 
Befaria racemosa Venten. (Ericaceae)—FLORIDA: St. Johns Co., St. Augustine, 17 
Jan. 1968, A. E. Graham, FSCA (ln). 


VOLUME 37, NUMBER 3 227 


Bumelia lanuginosa (Michx.) Pers. (Sapotaceae)—FLORIDA: Columbia Co., Iche- 
tucknee Springs St. Pk., em. 12 Jun. 1978, D. H. Habeck, FSCA (1n). 

Leucothoe populifolia (Lam.) Dipp. (Ericaceae)—FLORIDA: Marion Co., Juniper 
Springs, 30 Mar. 1979, D. H. Habeck, A-2349a, FSCA (1n). 

Vaccinium arboreum Marsh.—FLORIDA: Columbia Co., Santa Fe River, 8 mi. S Ft. 
White, c. 6 Apr. 1974, p. 9 Apr. 1974, em. 15 Apr. 1974, D. H. Habeck, FSCA (2n). 

V. stamineum L.—FLORIDA: Alachua Co., Gainesville, 30 Apr. 1972, em. 17 May 
1972, D. H. Habeck, FSCA (2n). 

Zomaria rosaochreana (Kearfott) 

Lyonia lucida (Lam.) C. Koch—FLORIDA: Glades Co., Palmdale, 18 Mar. 1973, D. 
H. Habeck, FSCA (8n); Alachua Co., Gainesville, 18 Aug. 1972, D. H. Habeck, FSCA 
(40n). 


LITERATURE CITED 


KARTESZ, J. T. & R. KARTESZ. 1980. A Synonymized Checklist of the Vascular Flora 
of the United States, Canada, and Greenland. Vol. II. The Biota of North America. 
Univ. of North Carolina Press, Chapel Hill. xlviii + 498 pp. 


Journal of the Lepidopterists’ Society 
37(3), 1988, 228-235 


NEW WISCONSIN BUTTERFLY RECORDS 


ROGER M. KUEHN 
546 Jordan Circle, Colgate, Wisconsin 53017 


ABSTRACT. Numerous new county records have increased the known range and 
relative abundance of many of Wisconsin’s butterflies (Rhopalocera). A number of species 
new to the state have been uncovered and several old records have been confirmed. The 
records were obtained from resident and non-resident collectors, published literature, 
university and natural history museum collections, and the author’s collection. 


Since the publication of Butterflies of Wisconsin (Ebner, 1970), I 
have maintained a list of all Wisconsin butterfly records. To date thir- 
teen new state records, six species of questionable occurrence and over 
1500 new county records have been added to this list. This brings the 
state total of confirmed butterfly species to 146. 

These records have come from much field work by resident collec- 
tors, various publications and field trips by myself. Collecting by George 
Balogh of St. Louis, Missouri, Robert J. Borth of Milwaukee, and Leslie 
A. Ferge of Middleton, particularly in prairies in the southern half of 
the state, in sphagnum-heath bogs in the north, and in oak-pine barren 
and bracken-grassland (‘‘stump prairie’) areas, has accounted for many 
new county records. Fay H. Karpuleon of Eau Claire and John H. 
Masters of California have added many new northern records. It should 
be noted that this intensive and widespread collecting has shown the 
ranges and relative abundance of many of Wisconsin’s butterflies to be 
much greater than is indicated in Butterflies of Wisconsin (Ebner, 
1970). 

In addition to the new state records, noted with a double asterisk 
(**) and confirmation of very old records, noted with an asterisk (*), 
only those new county records which are notable range extensions or 
relate to scarce or local species are mentioned in this supplement. 

Information regarding any Wisconsin butterfly records of which I 
am not aware would be greatly appreciated. 

All the species mentioned are single brooded in Wisconsin, except 
for those noted otherwise. A few species are also noted as representing 
strays due to their infrequent occurrence, undoubtedly from popula- 
tions to the south. 

The nomenclature and arrangement follow that of Ebner’s checklist 
(1970) but with some of the changes noted by Kuehn and Masters 
(1972). 


HESPERIIDAE 


**T erodea eufala (Edwards). Door, Douglas, Jefferson, Juneau, Milwaukee Counties, 5 
August-1 October. A small female in good condition was collected on 28 August 1966 
in Douglas County by Jackson L. Boughner and is now in the Milwaukee Public Museum 


VOLUME 87, NUMBER 3 229 


collection (Ms. Susan Borkin, pers. comm.). Since then two more specimens, also in good 
condition, were collected by George Balogh. The determination of these specimens was 
confirmed by Mogens C. Nielsen. Another specimen, a fresh female, was taken in Juneau 
County by Tom W. Kral. A fifth specimen was collected by William E. Sieker in Door 
County in the 1930's. This skipper could occur as a stray throughout Wisconsin, as it has 
been found as far north as the Upper Peninsula of Michigan (Nielsen, 1970). Because of 
its drab color and late flight period, eufala could be easily overlooked. It should be looked 
for in dry, open fields in fall. 


*Atrytonopsis hianna (Scudder). Adams, Burnett, Douglas, Eau Claire, Grant, Juneau, 
Wood Counties, 14 May—28 June. Found in the oak-pine barren areas of western and 
central Wisconsin, hianna flies with Lycaeides melissa samuelis Nabokov. Although 
widely distributed, it has not been taken in large numbers. This skipper was said to have 
been common in the Racine County area (southeastern Wisconsin) prior to the turn of 
the century (Hoy, 1883). There have been no subsequent records from this area. 


Euphyes conspicua (Edwards). 1 July-10 August. Twenty scattered new county records 
indicate that this skipper should be found statewide in sedgy meadow or marsh habitats 
and not only in those counties near Lake Michigan as suggested by Ebner (1970). In its 
habitat conspicua can be moderately common. 


Euphyes bimacula (Grote & Robinson). Barron, Chippewa, Douglas, Florence, Forest, 
Iowa, Juneau, Kenosha, Marinette, Racine, Vilas, Waukesha Counties, 25 June-30 July. 
These widely scattered new county records indicate a statewide distribution for bimacula. 
It is very local and never common in sedgy meadow or marsh habitats. 


Poanes viator (Edwards). 4 July—7 August. While generally scarce, this species has been 
found to be locally abundant at times in Dodge and Ozaukee Counties. 


**Problema byssus (Edwards). Grant County, 12 July 1981. Two males and a female, all 
fresh, were taken in prairie habitat in extreme southwestern Wisconsin by George Balogh 
and Robert J. Borth; another female was taken by James C. Parkinson in the same area 
on 11 July 1982. This species occurs in Iowa and throughout much of Illinois in areas on 
or near major rivers (Irwin & Downey, 1973). Thus it could also be expected in similar 
areas of southern Wisconsin. 


Atalopedes campestris (Boisduval). Brown, Dane, Eau Claire, Grant, Green, Jefferson, 
Juneau, Polk, Rock, Winnebago Counties, 12 August-20 October. New county records 
from Polk County in the northwest to Winnebago County in the east and Jefferson 
County in the south show that this skipper strays throughout the southern two-thirds of 
Wisconsin, especially during late summer and fall. 


Pompeius verna (Edwards). Brown, Dane, Douglas, Dunn, Grant, Iowa, Marathon, Mil- 
waukee Counties, 23 June-31 July. The distribution of verna may well be statewide 
based on these widely scattered new county records. 


*Polites origenes (Fabricius). Adams, Dane, Chippewa, Eau Claire, Grant, Green, Iowa, 
Juneau, Monroe, Pierce, Sauk, Waukesha Counties, 25 June—29 July. Records for origenes 
show it to be found at least through the southern half of Wisconsin. It flies in the same 
open, grassy fields as the similar and generally more common Polites themistocles (La- 
treille) with which, at first glance, it can easily be confused. This may be the reason that 
its presence in Wisconsin has gone unnoticed since the turn of the century (Rauterberg, 
1900). In dry prairie habitats, such as those found in Grant, Green and Sauk Counties, 
origenes often outnumbers themistocles. 


Hesperia metea Scudder. Adams, Brown, Douglas, Florence, Juneau, St. Croix Counties, 
12 May-1 June. The Adams and Juneau County records show that this skipper occurs as 
far south as central Wisconsin in oak-pine barren areas. It can be easily missed as the 
flight period is very short. 


Hesperia ottoe Edwards. Crawford, Dane, Grant, Green, Sauk, Waushara Counties, 24 
June-26 August. Found in modest numbers in the southwestern portion of the state, ottoe 
is restricted to dry prairie habitat. 


230 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Hesperia sassacus Harris. 15 May-29 July. Well over a dozen new county records show 
this skipper to be a resident of all but the most extreme southern counties of Wisconsin, 
having been found as far south as Iowa County. 


Hesperia leonardus Harris. Brown, Chippewa, Columbia, Dane, Douglas, Eau Claire, 
Florence, Green, Juneau, Langlade, Marathon, Oconto, Sauk, Shawano Counties, 4 Au- 
gust—10 September. While typical leonardus occur in northeastern Wisconsin, specimens 
from the western and southern portions of the state are part of a blend zone between 
leonardus to the east and Hesperia leonardus pawnee Dodge to the west (Dr. William 
W. McGuire, pers. comm.). The southernmost records (Green County) are of specimens 
showing varying degrees of transition between leonardus and pawnee; here they are 
found in dry prairie habitat. 


Hylephila phyleus (Drury). Dane, Grant, La Crosse, Manitowoc, Portage, Waukesha 
Counties, 2 August-15 October. These new county records indicate that phyleus strays 
throughout the southern half of Wisconsin as far north as Portage County (Johnson & 
Malick, 1972); it is seldom common. 


Thymelicus lineola (Ochsenheimer). Dane, Kenosha, Marinette, Ozaukee, Sheboygan, 
Vilas, Washington, Waukesha Counties, 16 June-21 July. This species is now common 
in the Milwaukee County and Dane County (Madison) areas and appears to be extending 
its range throughout eastern Wisconsin. 


*Oarisma poweshiek (Parker). Walworth, Waukesha Counties, 27 June-16 July. Two 
strong colonies of this skipper were discovered by George Balogh in low prairie areas on 
4 July 1978 in Waukesha County. Since then another locality may have been found when 
Leslie A. Ferge and James C. Parkinson, collecting in Walworth County, took a single 
specimen. Both of these counties are in southeastern Wisconsin. The last report of pow- 
eshiek in Wisconsin was by Fernekes (1906) in Milwaukee County. 


**Nastra lherminier (Latreille). Green County, 26 August 1979. A single female of this 
species was taken by George Balogh in a dry, upland prairie. This specimen was examined 
by me and, as /herminier has been reported from northcentral Minnesota (Azevedo, Jr., 
1970) and central Illinois (Irwin & Downey, 1973), it is likely this skipper will be found 
on occasion in Wisconsin. 


Carterocephalus palaemon mandan (Edwards). 20 May-5 July. Numerous new county 
records indicate that this species occurs in all but the southern quarter of the state, and 
it has been found as far south as Dodge and Washington Counties. 


Erynnis icelus (Scudder & Burgess). 6 May-8 July. Almost three dozen new county 
records show icelus to be common throughout Wisconsin. 


Erynnis brizo (Boisduval & Leconte). 24 April-14 June. Over a dozen new county records 
indicate that brizo is also found statewide, but it tends to be less common than icelus. 


*Erynnis persius (Scudder). Adams, Burnett, Eau Claire, Juneau, Monroe, Polk, Wood 
Counties, 11 May-—8 June. Not reported since before the turn of the century (Hoy, 1883), 
persius is seldom common in the oak-pine barren areas of central and western Wisconsin. 
It is associated with Lupinus perennis (blue lupine) and is found in company with 
Atrytonopsis hianna and Lycaeides melissa samuelis Nabokov (Lycaenidae). The Eau 
Claire County specimens were reared from ova found on L. perennis by Fay H. Kar- 
puleon and identified by Mogens C. Nielsen. Some of the Adams, Burnett and Wood 
County specimens were determined by Dr. John M. Burns. 


Erynnis lucilius (Scudder & Burgess). 9 May—25 September. Seventeen new county rec- 
ords for this skipper show it to be well distributed within the southern half of Wisconsin 
as far north as Chippewa County. Some late capture dates would indicate that a partial 
third brood occurs. 


Erynnis baptisiae (Forbes). Iowa, Iron, Juneau, Polk, Wood Counties, 3 July—5 Septem- 
ber. Robert P. Dana took several examples of this species in Polk County in 1973, and 


VOLUME 87, NUMBER 3 231 


also obtained some adults from larvae found on Astragalus canadensis (milk-vetch). The 
Iron and Wood County records and identification of the Polk County specimens were 
obtained from Dr. John M. Burns. A single specimen taken by George Balogh in Iowa 
County was determined by Richard Heitzman; a specimen also has been taken in Juneau 
County by Tom W. Kral. Dates for the few specimens taken in Wisconsin indicate a 
single brood. 


Erynnis martialis (Scudder). Burnett, Douglas, Eau Claire, Juneau, Waukesha Counties, 
15 May-18 August. These widely scattered records indicate that martialis may be found 
over much of southern and western Wisconsin. It is double brooded and common at 
times. 


Thorybes bathyllus (J. E. Smith). 6 June-19 July. Almost a dozen new county records 
indicate that this species is found throughout the southern half of Wisconsin. It had 
previously appeared to be limited to the southeastern portion of Wisconsin (Ebner, 1970). 


*Achalarus lyciades (Geyer). Buffalo County, 10 July 1956. Reported long ago by both 
Hoy (1883) and Rauterberg (1900), this single specimen is the only remaining known 
record from Wisconsin. Taken by Dr. John S. Nordin, he reported the species as being 
moderately common in clearings in wooded bluffs along the Mississippi River. 


PIERIDAE 


Artogeia virginiensis (Edwards). Florence, Langlade, Lincoln, Marathon, Oconto, Onei- 
da Counties, 4 May—2 June. Apparently confined to the northeastern quarter of Wiscon- 
sin, virginiensis occurs as far south as Marathon County. It is quite local and generally 
not too common in beech-maple forests. 


Colias interior Scudder. 4 June—31 August. This species occurs as far south in central 
Wisconsin as Juneau, Monroe and Waushara Counties. Fresh specimens taken on 19 
August 1971 and 22 August 1974 in Marathon County suggests that an occasional partial 
second brood occurs. 


Phoebis sennae eubule (Linnaeus). Grant County, 31 August 1975 (leg. Leslie A. Ferge), 
7 & 10 September 1931. Each date represents the capture of a single specimen. S. E. 
Ziemer (pers. comm.) reports that this species was regularly sighted in the 1930's in 
Kewaunee County. Since the middle 1950's, none has been seen there. 


Phoebis philea (Johansson). One perfect male was taken by S. E. Ziemer in Kewaunee 
County on 20 September 1930. More recently, a male in good condition was taken by 
Mrs. Kathleen Lukasavitz near Hartford, Washington County on 18 August 1979. This 
specimen was caught by using a small fruit basket in a flower garden! 


**Phoebis agarithe (Boisduval). Eau Claire County, 5 August 1979. A single, slightly 
worn male was taken by Fay H. Karpuleon in a gravel pit filled with grasses, weeds and 
wildflowers near the Chippewa River. As with the other members of the genus Phoebis 
mentioned, this specimen obviously represents a rare straggler in Wisconsin. 


Eurema mexicana (Boisduval). On 24 June 1977 a single male was collected in Sauk 
County. The specimen’s near perfect condition suggests that it may represent the off- 
spring of a migrant parent (James C. Parkinson, pers. comm.). There is also an example 
of this species from Wausau, Marathon County (no date) in the collection of Julia Wood 
(Russell A. Rahn, pers. comm. ). 


Falcapica midea annickae dos Passos & Klots. Waukesha County, 14-27 May. A single 
colony of this butterfly was located by Charles A. Kondor, Sr. in 1980 and eight specimens 
were taken. Additional specimens were observed and collected in 1981. Found only in 
very limited numbers, annickae occurs only in the immediate area of a small, dry, sandy 
wash with scattered oaks (Quercus sp.). 

As subsequent visits to the area by a number of collectors in 1982 and 1988 failed to 
locate additional specimens, the present status of this species in Wisconsin is uncertain. 


232 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


RIODINIDAE 


Calephelis muticum McAlpine. Dane, Fond du Lac, Sauk, Walworth, Waukesha Coun- 
ties, 11 July-17 August. This species should be found in the southern third of Wisconsin 
in marshes and wet prairies where Cirsium muticum (swamp thistle) occurs. It is at 
times moderately common, but extremely local. 


LYCAENIDAE 


**Satyrium caryaevorum (McDunnough). Adams, Dane, Douglas, Grant, Green, Green 
Lake, Iowa, Juneau, Lincoln, St. Croix, Waushara Counties, 27 June-2 August. These 
few records by several different collectors indicate that this hairstreak is found statewide 
but it is local and uncommon. Its similarity to the common Satyrium calanus falacer 
(Godart) undoubtedly has caused the misidentification of many specimens of caryaevo- 
rum. 


Incisalia polios Cook & Watson. Adams, Bayfield, Burnett, Douglas, Eau Claire, Flor- 
ence, Juneau, Marathon, Oneida, Washburn, Vilas Counties, 6-31 May. Based on these 
widely scattered new county records, this elfin should be found throughout the northern 
half of Wisconsin. It is often common but local on sandy, barren grasslands and oak-pine 
barrens. 


**Incisalia irus (Godart). Adams, Juneau Counties, 4-29 May. One female was collected 
by George Balogh on 21 May 1977; the determination was by Patrick J. Conway. Since 
then well over a dozen specimens have been taken here, in another nearby locality in 
Adams County and in adjacent Juneau County (leg. Robert J. Borth, Leslie A. Ferge, 
Tom W. Kral, Roger M. Kuehn and James C. Parkinson). The habitat is sandy, open 
woods with Lupinus perennis (blue lupine), typical of Wisconsin’s oak-pine barren areas. 


Incisalia henrici (Grote & Robinson). Chippewa, Juneau, Langlade, Oneida, St. Croix 
Counties, 1 May-6 June. Apparently quite local and not too common, henrici could be 
expected throughout northern Wisconsin, especially in the northwest quarter. In Burnett 
County it is moderately common in brushy, sandy oak-pine barren areas. 


Incisalia augustus (Kirby). 8 May-17 June. Over a dozen new county records show 
augustus to be found throughout the northern portion of the state and as far south as 
Adams County in central Wisconsin. It is fairly common throughout its range, especially 
in bogs. 


**Frora laeta (Edwards). Menominee County, 11 & 22 May 1968. Two specimens of this 
rare butterfly were reported by Richard A. Bailowitz (pers. comm.). As it has been taken 
in the Upper Peninsula of Michigan (Oosting, 1979), laeta should be looked for, but not 
necessarily expected, in Canadian Zone forest throughout northern Wisconsin. 


Epidemia dorcas (Kirby). Langlade, Lincoln, Marathon, Oneida, Rusk, Sawyer, Vilas 
Counties, 2 July-2 August. Distributed throughout much of the northern half of Wis- 
consin, dorcas is often moderately common but local. It occurs in and near marsh or bog 
habitats where Potentilla fruticosa (shrubby cinquefoil) is found. 


Epidemia epixanthe michiganensis (Rawson). 21 June-15 September. Thirteen new county 
records indicate that while michiganensis occurs mainly in northern Wisconsin, it is 
found as far south as Juneau, Monroe and Sheboygan Counties. It is at times very common 
in open bogs. 


*leptotes marina (Reakirt). Grant County, 16 July 1978. Two specimens were taken in 
extreme southwestern Wisconsin; their fresh condition made them appear to be newly 
emerged (Robert J. Borth, pers. comm.). These most likely represent the offspring of 
migrants, as this blue is seldom encountered to the south in Illinois (Irwin & Downey, 
1973). This butterfly had been reported from Milwaukee County many years ago (Fer- 
nekes, 1906). 


Hemiargus isola (Reakirt). Eau Claire, Grant, Price, Sawyer, Trempealeau Counties, 21 
June-21 August. These records indicate the sporadic occurrence of isola in western 
Wisconsin. 


VOLUME 37, NUMBER 3 233 


Lycaeides argyrognomon nabokovi Masters. 29 June—15 July. This species appears to be 
limited to the northeastern corner of the state with only Florence and Langlade Counties 
being new records. It is still present in the Waubee Lake area (George Balogh, pers. 
comm.) from which it was first reported by Louis Griewisch (1953). As Ebner (1970) 
was unaware that Lycaeides melissa samuelis occurred in Wisconsin, the true identity 
of specimens referred to from Brown and especially Burnett and Waupaca Counties is 
in doubt. L. a. nabokovi is common at times, but is extremely local in and near barren 
grassland openings on sandy soil in Canadian Zone forest. The foodplant in Wisconsin is 
Vaccinium caespitosum (dwarf bilberry) (Nielsen & Ferge, 1982). 


**T ycaeides melissa samuelis Nabokov. Adams, Burnett, Clark, Douglas, Eau Claire, 
Jackson, Juneau, Menominee, Monroe, Polk, Portage, Shawano, Waushara, Wood Coun- 
ties, 21 May—26 August. This blue is locally common in the oak-pine barren areas of 
central and western Wisconsin where its foodplant, Lupinus perennis (blue lupine), 
occurs (Masters & Karpuleon, 1975). There are two broods in Wisconsin. A superficial 
resemblance to L. a. nabokovi has probably led to the misidentification of specimens in 
the past. A female specimen from Portage County plated by Johnson and Malick (1972) 
as nabokovi is more likely referable to samuelis based on appearance and locality. 


**Everes amyntula ssp. (Boisduval). Burnett, Douglas, Washburn Counties, 22-28 May. 
Several examples of this western blue were taken on 27 May 1979 in sandy, oak-pine 
barren areas in Burnett County in extreme northwestern Wisconsin. Specimens of Everes 
comyntas (Godart) were also taken in the same areas. The identification of the Burnett 
County specimens (leg. George Balogh, Robert J. Borth & Leslie A. Ferge) was by Mogens 
C. Nielsen. As amyntula has been reported from central and northern Minnesota and 
the Upper Peninsula of Michigan (Mogens C. Nielsen, pers. comm.), it could possibly be 
found in Wisconsin’s northernmost counties. 


NYMPHALIDAE 


**Polygonia satyrus neomarsayas dos Passos. Douglas, Florence, Forest, Iron, Marathon, 
Marinette, Oneida, Sawyer, Vilas Counties, 2 June-3 September. These scattered records 
show that neomarsayas should be expected throughout northern Wisconsin. Found in 
company with Nymphalis vau-album j-album (Boisduval & Leconte) and Polygonia 
faunus (Edwards) in Canadian Zone forest, it has been taken in moderate numbers. 


Charidryas gorgone carlota (Reakirt). 15 May-15 September. Twenty-two new county 
records show carlota to be found statewide, except for the north central part of Wiscon- 
sin. Common at times, it is found in sandy, oak-pine barren areas in the western portion 
of the state and in dry prairies in southern Wisconsin. Fresh specimens taken in Septem- 
ber indicate that there are two broods in Wisconsin; the foodplant is Helianthus sp. 
(sunflower) (Wayne Duesterbeck, pers. comm. ). 


** Phyciodes pascoensis Wright. Florence, Fond du Lac, Forest, La Crosse, Marathon, 
Marinette, Oneida, Waukesha Counties, 11 June—5 August. As more resident collectors 
become aware of the presence of pascoensis and learn to distinguish it from the similar 
Phyciodes tharos (Drury), its range within Wisconsin will become much clearer. From 
the few reliable county records for pascoensis, it appears to occur in all but the southern 
quarter of Wisconsin. More attention will be required when collecting tharos and pas- 
coensis regarding flight dates, number of broods and type of habitat; their ranges may 
overlap in the Washington and Waukesha County areas. All determinations were made 
by Dr. Paul A. Opler. 


Phyciodes batesii (Reakirt). 30 May—2 August. Although found as far south as Adams 
and Juneau Counties, batesii is found mainly in the northern third of Wisconsin. It is 
occasionally found in moderate numbers in sandy, barren grassland habitat. 


Clossiana bellona (Fabricius). 8 May—29 September. Dozens of new county records show 
bellona to be found throughout Wisconsin, but it becomes less common and much more 
local southward in the state. Specimens from extreme northern Wisconsin are the sub- 
species Clossiana bellona toddi (Holland). 


234 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


**Clossiana frigga ssp. (Thunberg). Bayfield, Forest, Iron, Langlade, Lincoln, Oneida, 
Vilas Counties, 14 May-17 June. This species was discovered by four different collectors 
during the last week of May 1975 (Ferge & Kuehn, 1976). Although the records for 
frigga are concentrated in the north central portion of the state, it should be found 
throughout the northern third of Wisconsin in suitable bogs. Found in very wet, open, 
sedgy sphagnum-moss bogs, it is moderately common but the flight period is quite short. 


Clossiana freija ssp. (Thunberg). Bayfield, Langlade, Lincoln, Price, Vilas Counties, 10 
May-6 June. Restricted to the northern third of the state, freija is very local and generally 
uncommon in open sphagnum bogs. 


Proclossiana eunomia dawsoni (Barnes & McDunnough). Chippewa, Iron, Langlade, 
Lincoln, Marathon, Oneida, Price, Rusk, Sawyer, Vilas Counties, 23 May—27 June. First 
collected in a Price County bog by John H. Masters (Masters, 1971), dawsoni has been 
found in several north central Wisconsin counties. It is likely to be found throughout 
northern Wisconsin. 


HELICONIIDAE 


Agraulis vanillae nigrior Michener. Portage County, 12 May 1963. A second record of 
this species was reported from Wisconsin by Johnson and Malick (1972). There are also 
old reports from Milwaukee County (Muttkowski, 1907). 


SATYRIDAE 


**Satyrodes appalachia leeuwi (Gatrelle & Arbogast). Adams, Brown, Dane, Florence, 
Fond du Lac, Green Lake, Jefferson, Juneau, Manitowoc, Marathon, Marinette, Oconto, 
Ozaukee, Racine, Sauk, Walworth, Washington, Waukesha Counties, 23 June—21 August. 
This recently described subspecies (Gatrelle & Arbogast, 1974) has been found throughout 
southern Wisconsin and in scattered colonies in the northern portion of the state. As 
elsewhere, leeuwi occurs in both deciduous and tamarack swamps and forest-edge habitat 
adjacent to open marshes. Additional data is still required to determine the full extent 
of its range in Wisconsin. 


Coenonympha inornata Edwards. Ashland, Bayfield, Douglas, Iron Counties, 6 June-21 
July. At present, the range of this Ringlet appears to be limited to the extreme northwest 
corner of the state. It is found along roadsides and in open, grassy fields often in the late 
afternoon or early evening. 


Oeneis chryxus strigulosa McDunnough. Douglas, Florence, Langlade, Oneida, Vilas 
Counties, 14-31 May. These records indicate that strigulosa should be found throughout 
the northernmost third of Wisconsin. Found in jack pine barrens and bracken-grasslands 
(“stump prairies’), it is generally distributed and abundant at times. 


Oeneis jutta ascerta Masters & Sorensen. 14 May-—27 June. Eighteen new county records 
show ascerta to be concentrated in north central Wisconsin, although it has been found 
as far south as Monroe and Juneau Counties. It should be found throughout most of 
northern Wisconsin in black spruce/sphagnum-moss bogs. The more southern colonies 
in west central Wisconsin are in tamarack/sphagnum-moss bogs and represent the south- 
ern most records for this species in eastern North America. Ascerta is the most often 
encountered bog-related butterfly; the heaviest flights occur in odd-numbered years. 


Erebia discoidalis (Kirby). Iron, Langlade, Lincoln, Price, Vilas Counties, 14 May—6 June. 
As with many of northern Wisconsin’s bog-restricted species, discoidalis may or may not 
be present in any one particular bog. Found throughout the northern third of the state, 
it is very local but moderately common at times. 


VOLUME 37, NUMBER 3 235 


DISPOSITION OF SPECIMENS 


All specimens are presently retained by their respective collectors, except as noted 
below. 


Lerodea eufala leg. George Balogh, Roger M. Kuehn coll. (one specimen); leg. William 
E. Sieker, University of Wisconsin—Madison coll. 

Erynnis baptisiae leg. Robert P. Dana, National Museum of Natural History coll. (one 
specimen); Roger M. Kuehn coll. (one specimen). 

Achalarus lyciades leg. Dr. John S. Nordin, Roger M. Kuehn coll. 

Phoebis sennae eubule leg. unknown, University of Wisconsin—Platteville coll. (two spec- 
imens). 

Phoebis philea leg. Mrs. Kathleen Lukasavitz, Milwaukee Public Museum coll. 

Falcapica midea annickae leg. Charles A. Kondor, Sr., James A. Ebner coll. (two speci- 
mens); Philip A. Holzbauer coll. (two specimens); Roger M. Kuehn coll. (two speci- 
mens); Milwaukee Public Museum coll. (two specimens). 


ACKNOWLEDGMENTS 


I would like to thank all the persons who have made their collection records available 
to me and Mogens C. Nielsen and Dr. Allen M. Young who reviewed the manuscript. 


LITERATURE CITED 


AZEVEDO, JR., J. T. 1970. A Distributional Study of Butterflies in Itasca State Park, 
Minnesota. Mid-Cont. Lepid. Series, No. 18, 8 pp. 

EBNER, J. A. 1970. Butterflies of Wisconsin. Milwaukee Pub. Mus., Pop. Sci. Hndbk. 
No. 12, 205 pp. 

FERGE, L. A. & R. M. KUEHN. 1976. First records of Boloria frigga (Nymphalidae) in 
Wisconsin. J. Lepid. Soc. 30:233-234. 

FERNEKES, V. 1906. List of Lepidoptera occurring in Milwaukee County. Bull. Wisc. 
Nat. Hist. Soc. 4:39-58. 

GATRELLE, R. R. & R. T. ARBOGAST. 1974. A new subspecies of Lethe appalachia 
(Satyridae). J. Lepid. Soc. 28:359-363. 

GRIEWISCH, L. 1953. Lycaeides argyrognomon in Wisconsin. Lepid. News 7(2):54. 

Hoy, P. R. 1883. List of lepidopterous insects. Geol. of Wisc., 1873-1879, Vol. 1: 
23-25. 

IRWIN, R. R. & J. C. DOWNEY. 1973. Annotated checklist of the butterflies of Illinois. 
Ill. Nat. Hist. Survey, Biol. Notes No. 81, 60 pp. 

JOHNSON, K. & J. M. MaLick. 1972. An annotated list of central Wisconsin butterflies. 
Mus. of Nat. Hist., Univ. of Wisc.-Stevens Point, Rpt. No. 7, 6 pp. 

KUEHN, R. M. & J. H. Masters. 1972. Wisconsin butterflies. Mid-Cont. Lepid. Series, 
No. 59, 12 pp. 

MASTERS, J. H. 1971. First records of Boloria eunomia (Nymphalidae) in Wisconsin. 
J. Lepid. Soc. 25:149. 

Masters, J. H. & F. H. KARPULEON. 1975. Records of Lycaeides melissa samuelis 
(Lycaenidae) from Wisconsin. J. Lepid. Soc. 29:31. 

MuTTKowskKI, R. A. 1907. Additions to the lepidopterous fauna of Milwaukee County. 
Bull. Wisc. Nat. Hist. Soc. 5:128-133. 

NIELSEN, M. C. 1970. New Michigan butterfly records. J. Lepid. Soc. 24:42—47. 

NIELSEN, M. C. & L. A. FERGE. 1982. Observations of Lycaeides argyrognomon na- 
bokovi in the Great Lakes Region (Lycaenidae). J. Lepid. Soc. 36:233-234. 

OOsTING, D. P. 1979. Notes on the occurrence of Erora laeta (Lycaenidae) in Michi- 
gan’s western Upper Peninsula. J. Lepid. Soc. 33:149-150. 

RAUTERBERG, F. 1900. Diurnal Lepidoptera of Milwaukee County, Wisconsin. Bull. 
Wisc. Nat. Hist. Soc. 1:23-25. 


Journal of the Lepidopterists’ Society 
37(3), 1983, 286-243 


PUPAL COLOR DIMORPHISM IN CALIFORNIA 
BATTUS PHILENOR (L.) (PAPILIONIDAE): MORTALITY 
FACTORS AND SELECTIVE ADVANTAGE! 


S. R. Sims? AND A. M. SHAPIRO 


Department of Entomology, University of California at Davis, 
Davis, California 95616 


ABSTRACT. Estimates of Battus philenor (L.) pupal mortality were made in central 
California. Summer mortality of first and second generation pupae from unspecified 
causes ranged from 9-20%. Brachymeria ovata (Say) (Hymenoptera: Chalcididae) at- 
tacked and killed B. philenor in the pupal stage. Rates of parasitism varied between 
populations but not between pupae on narrow twigs or broad tree trunk habitats. 

A field experiment was conducted in a natural habitat of B. philenor to determine the 
selective advantage of pupal color dimorphism. Cryptic and non-cryptic pupae were 
affixed, in pairs, to narrow twigs in foliage or tree trunks and exposed to predators. Non- 
cryptic pupae in each pupation habitat suffered relatively more predation and lower 
survivorship. The extent of selective advantage conferred by cryptic coloration varied 
according to pupation substrate and season. Predation was greatest during the summer 
and on exposed tree trunks. The results indicate that B. philenor has greater survival on 
the pupation sites most frequently used in nature. 


The pupae of Battus philenor (L.) are dimorphic, being either green 
or brown with rare intermediates. In the central Appalachian Moun- 
tains of Virginia, philenor pupates off the ground on broad exposed 
surfaces such as tree trunks and cliffs (Hazel & West, 1979). These 
pupae are almost always brown. California philenor also pupate off 
the ground but much more frequently on narrow twigs within green 
foliage (Sims & Shapiro, 1983). Two-thirds (n = 1172) of the California 
pupae found on narrow twigs (<6 mm) are green. Pupae found on 
broad substrates such as grey concrete and brown tree trunks are pre- 
dominantly (92%, n = 283) brown. ; 

West and Hazel (1982) have shown higher survivorship of Virginia 
philenor pupae on the broad surfaces where they normally occur than 
on unutilized ground-level pupation sites in forest leaf litter. The dif- 
ference in mortality between the two types of sites was attributed to 
the relative palatability of pupae to different predators. Birds hunting 
above ground level probably learned to avoid the distasteful pupae; 
whereas, the greater palatability of pupae to small mammals hunting 
at ground level may have led to the formation of search images. 

The different pupation site distribution and color response of Cali- 
fornia philenor suggests that the selective mortality factors operating 
on apparent cryptic and non-cryptic pupae (background matching or 


1 Florida Agricultural Experiment Station Journal Series No. 5025. 
2 Current address: University of Florida AREC, 18905 S.W. 280th St., Homestead, Florida 33031. 


VOLUME 37, NUMBER 3 orl 


contrasting respectively) are distinct from those of the eastern United 
States populations (Sims & Shapiro, 1983). This is not surprising con- 
sidering differing factors in western areas such as the Mediterranean 
climate, riparian habitat of the foodplant (Aristolochia californica 
Torr.), evergreen nature of many associated dominant trees and shrubs, 
and probable differences in the species composition and predation pres- 
sure exerted by avian and other predators. 

Since present “non-preference” for certain available pupation sites 
is possibly due to continuing selection against individuals with this 
behavior (see Clarke & Sheppard, 1972 for evidence for a genetic basis 
of pupation site choice in Papilio polytes L.), we tested the hypotheses 
that 1) greater survival of both cryptic and non-cryptic pupae occurs 
on “preferred” twigs, and 2) cryptic pupae have a selective advantage 
on twigs as well as on broad tree trunk environments. 

Predation is not responsible for all mortality of philenor pupae. Pu- 
pae may be parasitized or die from undetermined causes. In this study 
we determined the percentage of pupae parasitized and compared 
rates of parasitization between pupae on narrow and wide substrates. 
We also estimated pupal death from undetermined causes over sum- 
mer and winter. 


METHODS AND MATERIALS 


We studied parasitization and other mortality patterns of philenor 
pupae at Goethe Park, Sacramento Co., CA (Latitude 38°40'N) (GP) 
and Bidwell Park, Chico, Butte Co., CA (39°45’N) (BP). Parasite data 
was also collected on larvae from the Vaca Mountains (from Mix Can- 
yon to Solano Lake, approximately 7 km SW Winters), Inner Coast 
Range, Solano Co., CA (38°25'N) (VM). 

B. philenor pupae from GP were collected on twigs and tree trunks 
in an area adjacent to a | km stretch of the American River on seven 
sample dates from 25 January to 5 March 1974. The vegetation at GP 
is an oak-dominated riparian forest. The most common tree and shrub 
species are live oak (Quercus agrifolia Nee.), elderberry (Sambucus 
mexicana Presl.), redbud (Cercis occidentalis Torr. ex Gray), coffee- 
berry (Rhamnus californica Esch.), and the larval host, Dutchman’s 
pipevine (Aristolochia californica). 

BP is also a riparian habitat dominated by oak. Pupae here were 
sampled from either twigs of the host plant growing on concrete sup- 
ports beneath a highway overpass or from the concrete itself. 

Mortality in some philenor pupae is manifested by external discol- 
oration and subsequent desiccation. This may result from disease, but 
the pathogen(s) remains undetermined. We estimated the magnitude 
of this type of mortality by collecting final instar larvae from VM and 


238 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


BP and allowing them to pupate and remain within outdoor cages in 
a sheltered location in Davis, Yolo Co., CA. 

We tested the selective advantage of cryptic pupal coloration on a 
0.5 km stretch of south-facing slope in Mix Canyon (VM population) 
at an elevation of 100 m. Both Aristolochia and philenor are common 
in this area. Diapause pupae used in this test had been field-collected 
on 11 June 1976 at BP. They were affixed to tree trunks and narrow 
twigs using clear silicone rubber sealant. On 8-9 July 1976, 64 pairs of 
green and brown pupae were attached to opposite sides of individual 
O. agrifolia tree trunks at heights of from 1 to 2 meters. An additional 
40 pairs were attached to narrow (<6 mm) twigs at similar heights in 
foliage of Q. agrifolia, Heteromeles arbutifolia M. Roem. (toyon), and 
Umbellularia californica (H. & A.) Nutt. (California bay). Pairs of 
pupae on twigs were separated by 30-40 cm. The location of pupae 
was marked by a stake at the base of the tree or shrub. Pupae were 
checked twice during the summer and once the following spring. 

West and Hazel (1982) recognized several possible fates for individ- 
ual pupae which are applicable here. A pupa may be: 1) alive and 
intact, 2) dead but intact, 3) attacked by a predator, remains visible, 
4) gone and presumably predated, 5) eclosed. We assume that missing - 
pupae are most likely predated and combine categories 3 and 4. The 
former position of pupae completely removed could generally be de- 
termined from the remaining traces of glue. The specific identities of 
probable predators were not determined in this study, although the 
visible remains of some predated pupae showed beak marks charac- 
teristic of birds. 


RESULTS 


The summer “disease” mortality, from June-September, of VM in- 
dividuals pupating during June in 1975 and 1976 was 20.3% (total 
n = 79) and 15.2% (n= 171), respectively. BP had 8.6% “disease” 
mortality (n = 558) during the summer of 1976. The incidence of the 
“disease’’ may be higher in summer than during the following fall- 
winter and spring. For example, a sample of BP pupae collected on 25 
January 1975 and monitored outdoors in Davis until emergence in 
March-April had only 4.6% (n = 195) dead pupae with these symp- 
toms. This may be variable from year to year. In warm and wet years 
(such as 1981-1982) fungal attack may be a significant cause of death 
in overwintering pupae. In later winter, pupae so attacked show mil- 
dew in their spiracles and intersegmental membranes, but this is rarely 
recognizable by late spring. 

During January-March 1974, 551 pupae were collected at GP. Most 


VOLUME 37, NUMBER 3 239 


pupae were found on stems and trunks of live oak, pipevine, redbud, 
and elderberry. Of 551 pupae, some appearing more than one year 
old, 427 (77.5%) were dead; 351 (82.5%) of the dead pupae had a 2- 
3 mm diameter circular opening in either the wing case or dorsum of 
thorax or abdomen suggesting a parasitoid emergence hole. Among 
dead individuals collected on narrow (<6 mm) twigs, 79% (n = 228) 
had “‘parasite” holes while 86% (n = 199) of dead pupae on broad tree 
trunks (=20 mm) had holes. These values are statistically similar (x2,, = 
3.54, P < 0.10). 

No parasitoids emerged the following spring from > 1000 diapausing 
winter-collected pupae (GP 1974; BP 1975, 1976), nor were any para- 
sitoids obtained from field-collected ova (>500 ova, VM 1973, 1974), 
3rd instar larvae (>100 larvae, VM 1974), or final instar larvae (>400 
larvae, VM 1974-1976; >200 larvae, BP 1975, 1976). 

Of 755 pupae collected 24 June 1975 at BP, 3.4% produced adult 
Brachymeria ovata (Say) (Hymenoptera: Chalcididae). All parasitoid 
emergence was completed by 12 July. On 11 June 1976, a sample of 
558 pupae had no parasitoids. Exit holes left by parasitoid adults from 
BP were identical to those seen in the GP pupae. B. ovata attacks and 
emerges from the pupal stage of many species of Lepidoptera (Har- 
ville, 1955; Peck, 1963). 

The fate of green and brown pupae on narrow twigs and broad 
trunks is shown in Table 1. We compared the survivorship of cryptic 
vs. non-cryptic pupae between sites by combining the number of suc- 
cessful adult emergences over the summer with the number of surviv- 
ing overwintering pupae. These comparisons are presented in Table 2. 
Most pupal mortality occurred during the summer, especially the first 
month following initiation of the experiment (Table 1). Green pupae 
on twigs and brown pupae on tree trunks suffered less predation than 
their alternate color forms but the differences were small (Table 2). 
The survivorship advantage of apparent cryptic coloration within sites 
in this study was largely due to greater adult emergence and lower 
non-predation mortality among the cryptic forms (Table 1). Our data 
suggests a lower survivorship of non-cryptic green pupae on tree trunks 
compared to brown pupae on twigs. Brown, and especially green pupae 
on tree trunks are quite conspicuous to the human observer and may 
be so to visually hunting predators. There was significantly reduced 
survivorship and increased predation on tree trunk pupae (combined 
green and brown) compared to twig sites (Table 2). 

We did not identify any of the predators of philenor, but the few 
remaining predated pupae had beak mark damage characteristic of 
birds. Most pupae were removed completely or with only a bit of the 
abdomen and cremaster remaining. 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


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240 


VOLUME 387, NUMBER 3 241 


TABLE 2. Survivorship (number of emerging adults during the year of the experi- 
ment + number of pupae alive the following spring) and predation of B. philenor pupae. 


Pupal color and substrate Survivors Predated x? (1) P 

Se eo on. tiga 24 ; 1.65 $0.10 

Fe on ce uunks 24 35 3.40 <0.10 

an Fie ai ee i o 45.28 <0.01 
DISCUSSION 


In this study we determined the magnitude of some philenor mor- 
tality factors in central California and the survivorship of cryptic and 
non-cryptic pupae on substrates which are most frequently used as 
natural pupation sites. 

An estimated 9-20% of intact pupae die from unknown, possibly 
pathogenic, agents during the summer months. One sample indicated 
that winter mortality of pupae with disease symptoms is somewhat 
lower than summer, but this may vary from year to year. 

The pupal-pupal parasitoid Brachymeria ovata attacks pupae in 
spring and summer. A single parasitoid is produced from each pupa, 
which is killed in the process. The parasitoids do not overwinter in 
diapause philenor pupae but probably overwinter as adults in a man- 
ner similar to other Brachymeria species (Clausen, 1940). Our deter- 
minations of percent parasitization may not accurately estimate the 
seasonal rate since we took only one sample each year per population. 
However, the rate of parasitization appeared low in the BP population, 
especially compared to the much higher estimate at GP. The deter- 
mination of annual percent parasitization at GP is complicated by our 
inability to determine what proportion of the dead pupae in our sam- 
ples represents individuals more than one year old. Thus, we have only 
an average estimate of parasitoid-caused annual mortality. 

It is possible that synchronization between parasitoid activity and 
the months of greatest philenor pupation (May-June) influences par- 
asitization rate. Individuals may not be “at risk’? during their entire 
pupation period, since Brachymeria prefers to oviposit in newly-formed 
pupae (Clausen, 1940). 

B. ovata has been observed to search randomly for host pupae in 
California habitats similar to those studied here (Harville, 1955). Ran- 
dom searching and host finding is reflected in the similar rates of 
parasitization of pupae on narrow and wide substrates. However, one 


242 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


of us (AMS) has observed a parasitization rate in excess of 90% in each 
of four years at a site (Rossmoor Bar, not far from GP) where a great 
many pupae are formed in the open on whitewashed fence posts; no 
data are available for pupae from more natural substrates. 

The primary pupal predators of California philenor remain unde- 
termined. Several local nocturnal animals such as the opossum (Didel- 
phis virginiana) and deer mouse (Peromyscus sp.) are possible candi- 
dates since they are often both arboreal and insectivorous (J. Harris, 
pers. comm.; Landry, 1970). Odor might be at least as important to 
these animals as visual cues. B. philenor adults are distasteful to some 
birds (Brower, 1958). Larvae are aposematically colored and it is likely 
that pupae possess alkaloids and aristolochic acid similar to those found 
in adults (Rothschild et al., 1970). We found some evidence for the 
unpalatability of pupae to birds since 5 of 82 pupae predated were 
clearly damaged by a bird’s beak but were left uneaten. Similarly, 
West and Hazel (1982) observed 18 of 139 damaged but uneaten phi- 
lenor pupae on tree trunks over a two year period in Virginia. 

The significantly reduced predation pressure among narrow pupa- 
tion substrates is correlated with the high percentage (83% of total) of 
brown pupae there. This suggests that brown pupae are as cryptic or 
otherwise protected as green pupae on twig sites and indicates the 
variability of the color-influencing stimuli of twigs. Tree trunks pro- 
duce less ambiguous color-determining cues. Only 8% of the pupae on 
broad exposed sites are non-cryptic green (Sims and Shapiro, 1983). 

Evidence is beginning to accumulate that a distinct selective advan- 
tage is accorded to individuals of dimorphic Lepidoptera species that 
choose an appropriate pupation site and have a cryptic pupal color 
response (Hidaka et al., 1959; Baker, 1970; Wiklund, 1975; West & 
Hazel, 1982). The choice of pupation site and ability of individuals to 
show a cryptic color response to the site’s color and texture varies in 
different philenor populations (West & Hazel, 1979; Sims & Shapiro, 
1983). We believe that this variability is related to a combination of 
the structure and seasonal phenology of the pupation-habitat coloration 
and the intensity of predation pressure. The latter is partly determined 
by predator species composition and density. In the deciduous forest 
of Virginia, most philenor pupae show a preference for rough exposed 
surfaces above ground level, are especially sensitive to the textural- 
optical qualities of these sites and, thus, pupate brown (West & Hazel, 
1979; Hazel & West, 1979). In the partly evergreen habitat determined 
by central California’s Mediterranean climate, narrow twigs in leafy 
areas are more frequently chosen, pupae are less sensitive to brown- 
producing stimuli, and more pupae are green (Sims & Shapiro, 1983). 
This study lends support to the hypothesis that pupation site preference 


VOLUME 387, NUMBER 3 243 


has evolved under differential selection by predators (West & Hazel, 
1982). The highest summer and overwinter survival and least predation 
occurred on the narrow pupation sites most frequently used by phile- 
nor in nature. 


ACKNOWLEDGMENTS 


We thank S. O. Mattoon for assistance in the study of the BP population, F. Gould 
and D. A. West for comments on an earlier draft of the manuscript, and C. Satterwhite 
for manuscript preparation. 


LITERATURE CITED 


BAKER, R. R. 1970. Bird predation as a selective pressure on the immature stages of 
the cabbage butterflies, Pieris rapae and P. brassicae. J. Zool. Lond. 162:43-59. 
BROWER, J. V. Z. 1958. Experimental studies of mimicry in some North American 
butterflies. Part II. Battus philenor and Papilio troilus, P. polyxenes and P. glaucus. 

Evolution 12:123-136. 

CLARKE, C. A. & P. M. SHEPPARD. 1972. Genetic and environmental factors influencing 
pupal color in the swallowtail butterflies Battus philenor (L.) and Papilio polytes L. 
J. Entomol. (A) 46:123-133. 

CLAUSEN, C. P. 1940. Entomophagous Insects. McGraw-Hill, NY. 688 pp. 

HARVILLE, J. P. 1955. Ecology and population dynamics of the California oak moth, 
Phryganidia californica Packard (Lepidoptera: Dioptidae). Microentomol. 20:83- 
166. 

HAZEL, W. N. & D. A. WEsT. 1979. Environmental control of pupal colour in swallow- 
tail butterflies (Lepidoptera: Papilioninae): Battus philenor (L.) and Papilio po- 
lyxenes Fabr. Ecol. Entomol. 4:393—400. 

HipAKA, T., T. KIMURA & M. ONOSAKA. 1959. Experiments on the protective coloration 
of pupae of the swallowtail, Papilio xuthus L. Zool. Mag. 68:222—226. (In Japanese 
with English summary.) 

LANDRY, S. O. 1970. The Rodentia as omnivores. Quart. Rev. Biol. 45:351-372. 

PECK, O. 1968. A catalogue of the Nearctic Chalcidoidea (Insecta: Hymenoptera). Can. 
Entomol. Supp. 30. 1092 pp. 

ROTHSCHILD, M., T. REICHSTEIN, J. VON Euw, R. APLIN & R. R. M. HARMAN. 1970. 
Toxic Lepidoptera. Toxicon 8:293-299. 

Sims, S. R. & A. M. SHAPIRO. 1983. Pupal color dimorphism in California Battus 
philenor (L.): Pupation sites, environmental control, and diapause linkage. Ecol. 
Entomol. 8:95-104. 

WEsT, D. A. & W. N. HAZEL. 1979. Natural pupation sites of swallowtail butterflies 
(Lepidoptera: Papilioninae): Papilio polyxenes Fabr., P. glaucus L. and Battus phi- 
lenor (L.). Ecol. Entomol. 4:387-392. 

1982. An experimental test of natural selection for pupation site in swallowtail 
butterflies. Evolution 36:152-159. 

WIKLUND, C. 1975. Pupal colour polymorphism and the survival in the field of cryptic 
versus non-cryptic pupae in Papilio machaon L. J. Roy. Soc. Lond. 127:73-84. 


Journal of the Lepidopterists’ Society 
37(3), 1983, 244-248 


A NEW SUBSPECIES OF SPEYERIA ATLANTIS 
(EDWARDS) (NYMPHALIDAE) FROM THE 
GREAT BASIN OF NEVADA 


GEORGE T. AUSTIN 


Nevada State Museum and Historical Society, 700 Twin Lakes Drive, 
Las Vegas, Nevada 89107 


ABSTRACT. A new subspecies of Speyeria atlantis (Edwards) of the unsilvered 
northern Great Basin cline from northeastern Nevada is described. This is the palest of 
the cline and occurs in an area known for pallidity in other Speyeria taxa. 


A number of new taxa of butterflies have been discovered and named 
from the more remote regions of the Great Basin over the past several 
years (Bauer, in Howe, 1975; Brown, 1975; Emmel & Emmel, 1971a, 
b; Emmel & Mattoon, 1972; Herlan, 1970; Howe, 1975; Scott, 1981; 
Shields, 1975). A distinct fritillary of the unsilvered, northern Great 
Basin cline of Speyeria atlantis (Edwards) has been known by a few 
collectors to occur in the Jarbidge and Independence ranges of north- 
eastern Nevada for about 20 years. The cline involved is largely unsil- 
vered, running from tetonia dos Passos and Grey from Wyoming 
through viola dos Passos and Grey in Idaho to dodgei (Gunder) in 
Oregon and ending in irene (Boisduval) in the Sierra Nevada of Cal- 
ifornia (see Moeck, 1957). The Nevada phenotype has usually been 
designated Speyeria atlantis near dodgei. It is, however, distinct enough 
to warrant recognition. 


Speyeria atlantis elko, new subspecies 


(Figs. 1 and 2) 


Description. Male, dorsal surface—Primaries and secondaries deep fulvous with the 
usual speyerian black markings moderately developed. Marginal band of primaries black 
with narrow lines of fulvous in each cell. Basal suffusion light to moderate on both wings. 
No lightening of ground color indicating positions of ventral surface pale markings. 
Ventral surface—Primaries basically pale tan with slight basal flush of fulvous. Markings 
of apical area and suffusion within marginal band a warm, slightly reddish, brown. On 
secondaries, all pale areas of same tan color as on primaries. Normal discal spots large 
and prominent. Narrower streaks of tan occur in median area of most or all cells. Disc 
pale brick red. Discal pale spots bordered basally with black; submarginal spots narrowly 
bordered distally with black. Basal spots in discal cell and cell Cu, usually completely 
encircled with black. Remaining dark areas of secondaries (marginal suffusion, basal 
border of submarginal spots) of same brown as markings of apical area of primaries. No 
silvering of any ventral spots. Size (all measurements of right primary along costal margin 
to furthest extent of apex)—Holotype = 27 mm, paratypes = 25-28 mm (N = 16). Ma- 
terial examined—Holotype and 28 paratypes. 

Female, dorsal surface—Ground color of primaries and secondaries of paler fulvous 
than male; black markings usually slightly less well developed. Marginal band of pri- 
maries tends to be filled completely with black apically but shows the fulvous ground, 
as in male posteriorly. Basal suffusion as in male. Ground color slightly lighter above 
positions of ventral pale spots. Ventral surface—Basic coloration similar to that of male 


VOLUME 387, NUMBER 3 245 


Fic. 1. Speyeria atlantis subspecies dorsal surface. Top row, males: left, elko, holo- 
type, NV: Elko Co.; Owyhee R. Valley, Wildhorse Creek Campgr., ca. 10 mi. S Mountain 
City, 8 July 1978, leg. G. T. Austin. Center, irene, CA: Nevada Co.; nr. Norden Lake, 
6700’, 26 July 1976, leg. C. Hageman. Right, dodgei, OR: Dead Indian Rd., 22 June 
1934, coll. unknown. Bottom row, females: left, elko, allotype, same data as holotype. 
Center, irene, CA: Nevada Co.; Soda Springs, 12 Aug. 1977, leg. B. O'Hara. Right, 
dodgei, CA: Siskiyou Co.; Methodist Camp, Castle Lake Rd., 27 July 1972, leg. L. P. 
Grey. 


but fulvous flush of primaries extending more into discal cell and noticeably to outer 
margin posteriorly. Pale spots of primaries and secondaries proportionally larger than 
those of males. Size—Allotype = 29 mm, paratypes = 28-30 mm (N = 10). Material ex- 
amined—Allotype and 15 paratypes. 

Types and type locality.* Holotype: NEV(ada): Elko Co(unty); Owyhee R(iver) Valley, 
Wild Horse Creek Campg(round), ca. 10 mi(les) S(outh) (of) Mountain City, 8 July 1978, 
leg. G. T. Austin. Paratype males: six with same data as holotype; two with same data 
except collected on 2 July 1980; one with same data except collected on 24 June 1981; 
six from Pine Creek, (Jarbidge Mountains) Elko Co(unty), Nevada, Jul(y) 10 (19)’72, leg. 
P. Herlan; three from Sawmill Creek, (Jarbidge Mountains) Elko Co(unty), Nevada, 
Jul(y) 8, (19)’74, leg. P. Herlan; one from same location, 7 (=July)-18-(19)76, leg. P. 


* Data on types are as indicated on specimen labels; parenthetical data correct errors or clarify label data; all from 
Nevada. 


246 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


7 


“e. 
aod 
oP 
>. 
> se 
y i 
Bs 3 


Fic. 2. Speyeria atlantis subspecies ventral surface. Same specimens as in Fig. 1 
(note especially the paleness and large spots of elko). 


Herlan; two from Elko Co(unty); Indep(endence) Range, Bull Run M(oun)t(ain)s, slope 
and summit of Porter Peak, 5 mi(les) W(est of) Maggie Summit, N(e)v(ada State Route) 
11A, 8000-9265’, 11 July 1982, leg. S. Mattoon; one from Elko Co(unty); Indep(endence) 
Range, N(e)v(ada State Route) 11A, Bull Run Basin to Columbia Basin at Aura Hist(orical) 
site, 6-10 mi(les) NNE H(igh)w(a)y 11, Deep Cr(eek) J(un)ct(ion), 6000-6300’, 11 July 
1982, leg. S. Mattoon; four from Elko Co(unty); Indep(endence) Range, Bull Run 
M(oun)t(ain)s, N(e)v(ada State Route) 11A, vic(inity) Maggie Summit, 6500’, 20 July 
1973, leg. S. Mattoon; two from Elko Co(unty); Indep(endence) Range, Bull Run 
M(oun)t(ain)s, N(e)v(ada State Routes) 226 and 1la, Jack Cr(eek) Campg(round) to Mag- 
gie Sum(mit), 5500-6619’, 10 July 1982, leg. S. Mattoon. 

Allotype: NEV(ada): Elko Co(unty); Owyhee R(iver) Valley, Wild Horse Creek 
Campg(round), ca. 10 mi(les) S(outh) (of) Mountain City, 8 July 1978, leg. G. T. Austin. 
Paratype females: two with same data as allotype; one from Jarbidge, (Jarbidge Moun- 
tains) Elko Co(unty), Nev(ada), Aug(ust) 11, (19)63, leg. P. Herlan; one from Jarbidge, 
(Jarbidge Mountains) Elko Co(unty), Nevada, Jul(y) 10, (19)’72, leg. P. Herlan; 11 from 
Elko Co(unty), Independence Range, Nev(ada State Route) 11A, 0.5 mi(les) E(ast) (of) 
Maggie Summit, 28 July 1981, leg. G. T. Austin. 

Due to the small number of specimens from any one specific location, the type series 
includes all specimens from Elko Co., Nevada, at hand. They were taken essentially from 
three colonies within 60 km of each other, one in the Jarbidge Mountains near Jarbidge, 
one in the Independence Range near Maggie Creek and the other along Wild Horse 
Creek, 6400’, in the Owyhee River Valley (R54E T44N S16). The latter was chosen as 
the type locality. The type locality is a creek bottom along which the males patrol. The 
surrounding area consists of low hills with sagebrush (Artemisia) as the predominant 


VOLUME 387, NUMBER 3 247 


vegetation. The new taxon flies with five other Speyeria: coronis snyderi (Skinner), 
zerene gunderi (Comstock), callippe harmonia dos Passos & Grey, egleis linda (dos Passos 
& Grey), mormonia artonis (Edwards). 

Deposition of types. The holotype, allotype, 10 male and seven female paratypes are 
deposited in the Nevada State Museum, one male paratype is deposited in the collection 
of C. S. Lawson in Las Vegas, Nevada, nine male paratypes are deposited in the collection 
of S. Mattoon, Chico, California, and the remaining paratypes are in the author’s personal 
collection. 

Other records. All NEVADA: Elko Co. (specimens not seen): Jarbidge Mts., Pine 
Creek, 9 July 1964 (J. Lane fide L. P. Grey); same location, 11 July 1972 (D. Bauer); 
same location, 10 Aug. 1967 (J. F. Emmel); Jarbidge Mts., Jarbidge-Charleston Road, 8 
mi. S. of Jarbidge, 9 Aug. 1967 (J. F. Emmel); Jarbidge River, 12 and 31 July 1974 (C. 
Ferris fide L. P. Grey); Rt. 11A, Maggie Creek, 21 July 1973 (L. P. Grey, Mattoon, fide 
L. P. Grey); same location, 21 July 1976 (L. P. Grey). 

Etymology. This subspecies is named after Elko County, Nevada, its type locality and 
only presently known range. 

Diagnosis. This new taxon is immediately recognizable from any other atlantis. 
The dorsal ground color is paler and the black patterning is finer than in tetonia, viola 
and dodgei. The males of those three subspecies tend to have the marginal area of the 
primaries largely black which is not the case in elko. In color, elko is similar to irene but 
the pattern is finer, especially on the secondaries, of elko. The ventral surface is partic- 
ularly distinctive. The reddish brown is paler in elko than in all the above-named taxa. 
The palest (aside from elko), irene, still shades towards a deeper brick red color which 
becomes progressively darker eastward. The submarginal band is wider, and the spots of 
the secondaries are larger than in any of the conspecifics, and the tan coloration of these 
gives the ventral surface of elko an almost yellowish appearance, an aspect not attained 
by any other taxon. 


Discussion 


Geographically, the paleness of elko corresponds closely with pallid- 
ity exhibited by other Speyeria of this same general region of the Great 
Basin (i.e., S. atlantis greyi Moeck, S. mormonia artonis, S. zerene 
gunderi). It is interesting that two very different clines of western 
atlantis coming from two directions terminate in extremes of pallidity 
within 80 km of each other. The populations of greyi in the Ruby 
Mountains and East Humboldt Range, Elko Co., Nevada may represent 
the western pallid extreme of the chitone (Edwards) and wasatchia 
dos Passos and Grey cline, while the taxon described herein is the pallid 
extreme of the northern Great Basin-Sierra Nevada cline which ap- 
parently colonized Nevada from the north. This situation approaches 
that shown by another pair of atlantis subspecies, hollandi (Chermock 
& Chermock) and dennisi (Gunder), which fly together in the Black 
Hills (Grey et al., 1963) and by a pair of S. zerene (Boisduval) subspe- 
cies, zerene and gunderi, which overlap in northeastern California 
(Grey & Moeck, 1962). 


ACKNOWLEDGMENTS 


Sincere thanks are due L. P. Grey for sharing his vast knowledge of Speyeria and 
useful comments for improvement of this paper. Thanks also to D. L. Bauer and J. F. 
Emmel for allowing use of their field data and S. O. Mattoon for the loan of specimens. 
Appreciation is due to Pam Church for her able typing and editing of the manuscript. 


248 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


LITERATURE CITED 


Brown, F. M. 1975. A new subspecies of Glaucopsyche (Phaedrotes) piasus from 
Nevada (Lepidoptera:Lycaenidae). Proc. Entomol. Soc. Wash. 77:501-504. 

EMMEL, T. C. & J. F. EMMEL. 197la. An extraordinary new subspecies of Cercyonis 
oetus from central Nevada (Lepidoptera, Satyridae). Pan-Pac. Entomol. 47:155-157. 

1971b. A new subspecies of Papilio indra from central Nevada (Lepidoptera: 
Papilionidae). Pan-Pac. Entomol. 47:220-223. 

EMMEL, T. C. & S.O. MATTOON. 1972. Cereyonis pegala blanca, a “missing type” in 
the evolution of the genus Cercyonis (Satyridae). J. Lepid. Soc. 26:140-149. 

Grey, L. P. & A. H. MOEcK. 1962. Notes on overlapping subspecies. I. An example in 
Speyeria zerene. J. Lepid. Soc. 16:81-97. 
GREY, L. P., A. H. MOECK & W. H. Evans. 1963. Notes on overlapping subspecies. II. 
Speyeria atlantis of the Black Hills (Nymphalidae). J. Lepid. Soc. 17:129-147. 
HERLAN, P. J. 1970. A new subspecies of Limenitis archippus (Nymphalidae). J. Res. 
Lepid. 9:217-222. 

Howe, W. H. 1976. The Butterflies of North America. Doubleday, Garden City, NY. 

MoEck, A. H. 1957. Geographic variability in Speyeria. Comments, records and de- 
scription of a new subspecies. Milwaukee Entomol. Soc., Special Paper. 

ScoTT, J. A. 1981. New Papilionoidea and Hesperioidea from North America. Papilio 
(new series), no. 1. 

SHIELDS, O. 1975. Studies on North American Philotes (Lycaenidae). IV. Taxonomic 
and biological notes, and new subspecies. Bull. Allyn Mus., no. 28. 


Journal of the Lepidopterists’ Society 
37(3), 1983, 249-252 


THE LARVA OF SIDERIDIS MARYX (GUENEE) 
(NOCTUIDAE) 


KENNETH A. NEIL 


Department of Biological Sciences, Simon Fraser University, 
Burnaby, British Columbia, Canada V5A 1S6 


ABSTRACT. The mature larva of Sideridis maryx (Guenée) (Noctuidae) is described 
and illustrated. 


The noctuid genus Sideridis (Hiibner) (Hadeninae) is represented in 
eastern North America by three species, S. rosea (Harvey), S. conger- 
mana (Morrison), and S. maryx (Guenée) (Forbes, 1954). Of the three 
species, S. rosea is the most common, and the immature stages are best 
known; they were most recently described and illustrated by Godfrey 
(1972) who listed grass, dandelion, Elaeagnus angustifolia L., Ribes 
sp., Shepherdia sp., and Salix sp. as host plants. S. congermana and S. 
maryx are generally considered to be uncommon to rare throughout 
eastern North America (Forbes, 1954; Rockburne & Lafontaine, 1976). 
Dyar (1899) briefly described the mature larva of S. maryx (=Ma- 
mestra rubefacta Morrison) based on preserved material. The larva of 
S. congermana and the natural host plants of all three species of Sid- 
eridis are unknown. 

This paper describes the mature larvae of S. maryx reared from ova 
obtained from a female taken on 11 June 1979 at Belliveau Cove, 
Digby Co., Nova Scotia. Larvae were fed an artificial diet based on 
that of Hinks and Byers (1976). They grew quickly and pupated by 7 
August. Adults emerged 10-15 September 1979. Throughout its range, 
S. maryx is single brooded, overwintering as a pupa, with adults emerg- 
ing in late spring and early summer. 

The terminology and abbreviations used here follow Godfrey (1972). 
The illustrations which accompany the description of the last larval 
instar were drawn to scale using a camera lucida and stereomicroscope. 


Sideridis maryx (Guenée) 


General. Head 3.5-4.0 mm wide. Total length 40.0-49.5 mm. Head and body smooth. 
Prolegs present on Ab3-6, size increasing posteriorly on Ab3-6, those on Ab3 slightly 
more than '% the size of those on Ab6. Crochets uniordinal, 19-22 per third abdominal 
proleg, 21-25 per fourth, 26-29 per fifth, 27-30 per sixth. All simple. 

Coloration (living material). Head (Fig. 3): yellowish brown with darker reddish brown 
coronal reticulations and coronal stripes. Body (Figs. 1, 2): red; dorsal and subdorsal areas 
with numerous greyish flecks, flecks heaviest at edges of middorsal line; middorsal and 
subdorsal lines narrow and poorly defined, the middorsal line more well defined on 
T1-3; lateral and ventral areas translucent red, lacking greyish flecks. Spiracles yellowish 
brown with black peritremes. Lateral shields of prolegs and thoracic legs yellowish brown, 
both darker basally. Prothoracic shield orange brown. 


250 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Fics. 1 & 2. Sideridis maryx, sixth instar larva (<8): 1, dorsal view; 2, lateral view. 


Head (Fig. 3). Epicranial suture 0.92-1.05 mm long; height of frons (apex to Fa’s) 
1.03-1.05 mm; distance from F1 to anterior edge of clypeus 0.33-0.85 mm; interspace 
between F1—-F1 0.45-0.47 mm; aFa anterior and Af2 posterior to apex of frons; Al-A3 
forming an obtuse angle at A2; P1-P1 1.54-1.56 mm; P2—P2 1.70-1.72 mm. Distance 
from P1 to epicranial suture less than that from P1 to L; L cephalad of juncture of 
adfrontal ecdysial line. Ocellar spacing: Ocl—Oc2 0.07-0.09 mm; Oc2—Oc3 0.10-0.12 
mm; Oc3-—Oc4 0.05-0.06 mm. 

Mouthparts. Hypopharyngeal complex (Fig. 4): spinneret short and broad, subequal 
to Lpsl; Lpsl longer than Lp2; stipular setae short, about 4 the length of Lps1, slightly 
shorter than Lpl, and about % the length of Lp2; Lps2 subequal to Lp1; distal and 
proximal regions of hypopharyngeal complex separated by a distinct medial transverse 
cleft; distal region with distal % bare, remainder with short thin spines becoming longer 
and slightly more robust proximally; proximolateral region with 15-20 stout spines. Man- 
dible (Figs. 5 & 6): two well-separated outer setae present; inner surface with ridges and 
tooth; inner tooth prominent, base broad, apex truncate; first outer tooth well developed, 
serrated on outer side; second outer tooth serrated on side opposite outer tooth; third and 
fourth outer teeth acutely angular; fifth outer tooth wide and flat with outer margin 
serrated. 

Thorax. Segment T1 (Fig. 8): prothoracic shield weakly sclerotized; SD1 and SD2 setal 
insertations well separated from shield; interspace D1—D1 about 0.65 XD1-XD1; D2- 
SD-2 about 1.41 SD2-—XD2; spiracle elliptical, 0.44-0.48 mm high, 0.27-0.29 mm wide; 


VOLUME 37, NUMBER 3 251 


ff ee  y. 


\' ; : me a We 
4 Ue eek 
[sf Ab7 Ab8 / x 


AD ae Ab10 


Fics. 8-8. Sideridis maryx, larva: 3, frontal view of head capsule; 4, left lateral view 
of hypopharyngeal complex; 5, oral surface of left mandible; 6, outer surface of left 
mandible; 7, dorsal view of anal shield; 8, dorsolateral chaetotaxy of prothoracic (T1), 
mesothoracic (T2), and abdominal segments (Ab1—-2, Ab6-10). Scale lines equal 1.0 mm. 


peritreme wider laterally. T2 (Fig. 8): Dl1-D2 about 0.69 D2-SD2; all setae thin and 
hairlike, tapering and sharply pointed distally; coxal bases narrowly separated. 

Abdomen. Dorsal and lateral chaetotaxy of Ab1-10 as in Fig. 8. Ab] with 2 SV setae, 
Ab2-6 with 3 SV setae, Ab7-8 with 1. Ab9: SD1 much finer than D1 and D2. Ab1O: 
Anal shield as in Fig. 7. Dorsal margin convex, posterior margin entire. Length of D1 
on Ab6-7 0.49-0.53 mm; D2 0.56-0.60 mm. Asp7 0.36-0.37 mm high, 0.21 mm wide; 
Asp8 0.49-0.50 mm high, 0.27 mm wide. 

Material examined. 4 specimens: Belliveau Cove, Digby Co., Nova Scotia. Reared on 
artificial diet (Hinks & Byers, 1976) from ova obtained from a female taken on 11 June 
1979. Adults emerged 10-15 September 1979. Moth collected, determined, and larvae 
reared by K. A. Neil. 


252 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Remarks. Based on the larval description and illustrations of S. rosea given by Godfrey 
(1972), S. maryx can easily be distinguished from that species by the truncate inner 
mandibular tooth, the short, stout spinneret which is only about % the length of the 
spinneret of S. rosea, and by the wider F1—F1 interspace. 


ACKNOWLEDGMENTS 


I thank Dr. G. L. Godfrey of the Illinois Natural History Survey for reviewing this 
manuscript and Ronald Long of Simon Fraser University for photographing the illustra- 
tions. 


LITERATURE CITED 


Dyar, H. G. 1899. Descriptions of the larvae of fifty North American Noctuidae. Wash. 
Entomol. Soc. Proc. 4:315-332. 

FORBES, W. T. M. 1954. Lepidoptera of New York and neighboring states. Pt. III. 
Cornell Univ. Agr. Expt. Sta. Mem. 329. 433 pp. 

GopFREY, G. L. 1972. A review and reclassification of larvae of the subfamily Had- 
eninae (Lepidoptera, Noctuidae) of America north of Mexico. U.S. Dept. Agr. Tech. 
Bull. 1450. 265 pp. 

HINKs, C. F. & J. R. BYERS. 1976. Biosystematics of the genus Euxoa (Lepidoptera: 
Noctuidae). V. Rearing procedures and life cycles of 36 species. Can. Entomol. 108: 
1345-1357. 

ROCKBURNE, E. W. & J. D. LAFONTAINE. 1976. The cutworm moths of Ontario and 
Quebec. Can. Dept. Agr. Publ. 1593. 164 pp. 


Journal of the Lepidopterists’ Society 
37(3), 1983, 253-254 


GENERAL NOTES 


COLLECTION RECORDS OF BUTTERFLIES FROM 
NYANGEZI, ZAIRE, AFRICA? 


Beginning in 1977, while working in the Peace Corps as a science teacher, I studied 
and collected butterflies from the small village of Nyangezi, in eastern Zaire, Africa, for 
approximately 20 months. 

Zaire is centrally located in Africa. It is bordered by the People’s Democratic Republic 
of the Congo on the west, Central African Republic and Sudan on the north, by Uganda, 
Rwanda, Burundi and Tanzania on the east and on the south by Zambia and Angola. 
The province of Kivu is in the eastern portion of the country, and the village of Nyangezi 
is in the southern third of Kivu, along the Rwanda and Burundi borders. 

Nyangezi is in the midst of montane highlands having an altitude of approximately 
1600-2000 m. It is partly located in a large interpluvial forest and has a very moist 
climate. The combination of the climate and altitude produces a constant temperature 
of approximately 24°C. The region was once richly endowed with native tropical flora 
but today introduced cypress, eucalyptus and various citrus species, as well as many 
agricultural plants, predominate. The village of Nyangezi would thus be termed a dis- 
turbed site for collecting butterflies. 

The families of Lepidoptera found in the village included: Hesperiidae, Papilionidae, 
Pieridae, Danaidae, Nymphalidae, Acraeidae and Satyridae. Unlike Rogers and van 
Someren (1925, E. Afr. Nat. Hist. Soc. J. 6:22—43), who treated these taxa as subfamilies, 
I assigned family rank, following the precedence of Williams (1969, A Field Guide to 
the Butterflies of Africa, Collins, London). The following species, arranged by family, 
were collected. 


HESPERIIDAE 


I collected only one specimen of Calaenorrhinus galenus (Fab.) although I observed 
others. 


PAPILIONIDAE 


Graphium simoni Aurivillius, Papilio cynorta Fabricius, P. dardanus Brown, P. de- 
modoceus Esper, P. echerioides Trimen, P. nireus L., P. zenobia and P. zoroastres 
Druce. 


PIERIDAE 


Belenois aurota Fabricius, B. zochalia Boisduval, Catopsilia florella Fabricius, C. 
thauruma (Reak.), Eurema brigitta Cramer, Nepheronia thalassina Boisduval. 


DANAIDAE 


Amauris albimaculata Butler, A. echeria Stoll, A. niavius L., A. ochlea Boisduval, 
Danaus chrysippus L., D. limniace Cramer. 


NYMPHALIDAE 


Byblia acheloia Wallengren, Cataeroptera cloanthe Cramer, Charaxes fulvescens Au- 
rivillius, Cymothe theobene Doubleday, Euphaedra spatiosa Mabille, Eurytela dryope 
(Cramer), Hamanumida daedalus Fabricius, Hypolimnas dinarcha Hewitson, H. mis- 
ippus L., Neptidopsia ophione (Cramer), Neptis nemetes Hewitson, N. saclava Bois- 
duval, N. seeldrayersi Aurivillius, Phalanta phalantha Drury, Precis hierta Fabricius, P. 
natalica Felder, P. octavia octivia Cramer, P. oenone L., P. sophia sophia Fabricius, P. 
terea terea Drury, Pseudacraea eurytus L., Pseudargumnis hegemone Godart, Salamis 
anacardi Trimen, S. parhassus Drury. 


‘ Approved by the Director of the North Dakota Agricultural Experiment Station as Journal Paper No. 1298. 


254 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


ACRAEIDAE 


Acraea asbolophintha (Karsh.), A. egina Mabille, A. woui Grose-Smith, Bematistes 
macarioides Aurivillius, B. poggei Dewitz. 


SATYRIDAE 


Mychalesis chapini Holland, M. langi Holland, M. saussure Dewitz, Ypthima albida 
Butler, Y. doleta Kirby. 


WILLIAM GLUCK, Graduate Research Assistant, Entomology Department, North Da- 
kota State University, Fargo, North Dakota 58105. 


Journal of the Lepidopterists’ Society 
37(3), 1983, 254-256 


NOTES ON THE GENUS IMELDA (RIODININAE) 


In “Illustrations of the Diurnal Lepidoptera,” volume 5(3), Hewitson described the 
riodinid butterfly Nymphidium mycea from a female received from “New Granada,” 
an area covering present-day Colombia, Venezuela, and Panama. His description reads 
as follows: “Upperside pale yellow, with the margins broadly dark brown; the outer 
margin of both wings transversed by a rufous band. Anterior wing with three white spots 
near the apex.” 

In 1879, Hewitson described a second butterfly based on a male from Ashpiyaco, 
Ecuador, which he named Imelda glaucosmia and designated it as the type of the genus 
Imelda, which was described in the same article. He later illustrated it in “Illustrations,” 
volume 4(5), plate 24, figure 5, repeating the original description, as follows: “Upperside 
of male glossy dark blue, slightly tinged with green. Both wings crossed beyond the 
middle by a narrow linear black band scarcely visible on the anterior wing; both with a 
submarginal band and the outer margin (which is broad) black. Anterior wing with the 
costal margin brown; crossed by a subapical broad band of white bordered with black. 
Underside as above, except that it is grey-brown, that each wing has two subbasal spots 
and a linear spot at the end of the cell of dark brown, and that the inner black band is 
much broader.” 

Thieme (1907, Berlin Ent. Zeitschrift 52:1-16) designated mycea as the female of 
glaucosmia, using the name mycea to refer to glaucosmia specimens from Colombia. 
H. Stichel (1910, Berlin Ent. Zeitschrift 55:9-103) erected a new subspecies, terpna, to 
refer to the male designated by Thieme as that of mycea. Stichel rejected the idea that 
mycea was the female of terpna, claiming that the dimorphism between the two was 
too great for them to be conspecific. He maintained this position in the Catalogus (Stichel, 
1930, in Junk, Lepidoptorum Catalogus, Vol. 44, Berlin). 

Because the most recent revision of a taxonomic group usually takes precedence over 
previous revisions, Stichel’s conclusions determine the present status of these butterflies. 
However, as both Thieme’s and Stichel’s conclusions were reached without the help of 
field observations, the matter is worth reopening in light of data I gathered in the field. 

My first experience with glaucosmia came during a collecting trip to a locality about 
14 km to the west of Arcabuco, Boyaca, Colombia, in July 1981. The altitude of this area 
is 2000 m and lies in a transition zone between Very Humid Low Montane Forest and 
Premontane Very Humid Forest. Rainfall is about 2000 mm per year (1977, Anonymous, 
Zonas vegetales de Colombia, IGAC). Although much of the vegetation has been cleared 
for cattle raising, forested areas may be found along the streams. The general aspect of 
the forest is like other subtropical montane forest areas throughout the neotropical region. 
The trees reach a height of 10 m and support many bromeliads and other epiphytic 
plants growing from the branches, and are interspersed with bamboos and tree ferns. 


VOLUME 87, NUMBER 3 255 


Fics. A-H. Imelda mycea ssp. (dorsal and ventral views): A & B, male I. m. mycea 
Hewitson, 6 & 2 from Arcabuco, Boyaca, 2000 m, Colombia; € & D, female I. m. mycea; 
E & F, male I. m. glaucosmia Hewitson, é & ? from Rio Topo, 1500 m, Tungurahua, 
Ecuador; G & H, female I. m. glaucosmia. 


256 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


At 1245 h while collecting in a small forest clearing by a stream, I noticed a small 
blue riodinid butterfly resting beneath a leaf with wings outspread at the edge of the 
clearing about 3 m above the ground. I had barely captured this butterfly when another 
indentical individual alighted on the same spot. This butterfly was also caught. Moving 
along the edge of the clearing, I captured another riodinid butterfly, this time a yellow 
one, resting under another leaf about 4 m off the ground. All this took place within a 5 
minute period. The behavior exhibited by these butterflies was typical of perching, a 
form of mate locating behavior employed by members of the subfamily Riodininae, in 
which the butterflies wait for mates at certain localities and during certain hours of the 
day (Callaghan, in prep.). 

During a subsequent visit to the same locality on 16 September 1982, four additional 
males and one female were captured between the hours of 11386 and 1320. All were 
frequenting the same microhabitat and exhibiting the same behavior as on the previous 
visit. 

Through consulting the descriptions and references above, I determined the butterflies 
to be Imelda glaucosmia terpna Stichel and Imelda mycea Hewitson. 

In view of the behavior observed and the morphology of the butterflies, I conclude 
that terpna and mycea are male and female of the same species. First, my studies of the 
perching habits of riodinid butterflies have shown that the frequenting of similar perch- 
ing sites at the same time by closely related male and female phenotypes is a strong 
indication that the two are conspecific. Secondly, there is enough similarity in the mor- 
phology of glaucosmia terpna and mycea to suggest that they are conspecific. The 
general pattern with the white spots on the apex of the forewing and the placement of 
the submarginal bands on both, as well as the marginal row of white spots on the 
underside of both wings is sufficient indication that the two are indeed conspecific as 
indicated by Thieme. 

In conclusion, the name Imelda mycea mycea (Hewitson, 1865, [1852-1878], Illustra- 
tions of Diurnal Lepidoptera, vol. 1-5) refers to central Colombian material, illustrated 
in Figs. A, B, C and D, for which the name terpna Stichel is a synonym. The name 
Imelda mycea glaucosmia (Hewitson, 1870, Ecuatorial Lepidoptera, Part IV) is the 
designation for material from Ecuador to Southern Colombia (Huila), illustrated in Figs. 
E, F, G and H. 

I wish to thank Dr. Keith Brown for his comments on the paper. | 


CurtTIs J. CALLAGHAN, © Pesquisador Associado,” Museu Nacional, Rio de Janeiro, 
Brazil. 


Journal of the Lepidopterists’ Society 
37(3), 1983, 256-257 


NEW AND UNUSUAL BUTTERFLY RECORDS FROM KANSAS 


In June 1979, my colleagues and I (senior author) began an intensive, statewide survey 
of the butterflies of Kansas. Our third season (1981) was marked by an influx of immigrant 
species and the collecting of two species (Pyrisitia proterpia and Speyeria edwardsit) not 
previously reported from Kansas. Observations and collections were also made for Thes- 
salia fulvia and Vanessa annabella, both considered “unusual” for Kansas. 

Pyrisitia proterpia (Fabricius).—Labedz took a single female (nearly perfect condi- 
tion) on the Fort Hays State University campus, Hays, Ellis County on 20 October 1981. 
This individual was taken on a purple-flowered, ornamental Amaranthus sp. at about 
1445 h. The temperature was about 70°F with a light wind and the sky was clear. Upon 
seeing this specimen, Rolfs recalled collecting a similar one on 3 October and of seeing 


VOLUME 87, NUMBER 3 WAS) 


a second “a few days later’ (about 7 October). These two individuals were over a 
vegetable and flower garden in north Hays. No additional specimens were found despite 
an extensive search on following days. William Howe (pers. comm.) captured a tattered 
vagrant in Franklin County in October 1971 (now at LACM). These are the first reported 
specimens from Kansas. 

Speyeria edwardsii (Reakirt).—Ely collected a male in perfect condition in Atwood, 
Rawlins County on 5 June 1981. It was feeding in a small patch of alfalfa (Medicago 
sativa L.) growing at the edge of a paved street at the edge of town. Nine other species 
were collected at this site, and we spent about 30 minutes searching, in vain, for another 
specimen. The weather was sunny and warm (late afternoon) with a light wind. This is 
the first reported Kansas specimen. The only other unusual species seen in Rawlins 
County on this date was a single Danaus gilippus which narrowly evaded capture at 
another locality. 

Thessalia fulvia (W. H. Edwards).—Field (1938, A manual of the butterflies and 
skippers of Kansas, Bull. Univ. Kansas 39:328 pp.) reported a specimen from Rush County 
on 28 June 1912. Marvin D. Schwilling (pers. comm.) took three specimens (now in his 
private collection) in Barton County during the 1960's. Ely took a fresh but slightly 
damaged specimen from a truck radiator on 14 June 1980. This specimen must have 
been hit between Holcomb (Finney County) and Hays and we suspect closer to the latter. 
This was followed by multiple finds by Ely in Ellis County. 

On 15 June while on a class field trip to the FHSU pasture about one mile south of 
Hays, he collected a single male (by hand) in moderately grazed mixed grass prairie. 
Next morning on a similar trip he saw several others in a nearby area, so returned after 
class for a more extensive search. Between 1020 h and 1100 h, seven of the 15 individuals 
seen along a white chalk road through the prairie were collected. They appeared to be 
attracted to the road since only three were found on a similar intersecting road nearby 
and none was seen in adjacent prairie. All seemed to be males and some seemed “terri- 
torial” in that each repeatedly returned to its same spot after being flushed. All alighted 
on the ground rather than on vegetation and no feeding was observed. The area was 
rich in flowering forbs and pioneer plants. The weather was warm and without wind. 

Ely next visited the area on the 17th (1400-1420 h), a warmer day with moderate 
wind, and failed to find even one individual. A careful examination of the Castaleja 
plants in the vicinity failed to locate any larvae. On the 24th he again visited the area 
during mid-morning (1000-1010 h) and saw at least 10 individuals, including one feeding 
on Houstonia nigricans (Lan.) Fern. The area was last visited on 4 July but no fulvia 
were found. During this entire period all individuals were within an area of approxi- 
mately 10 m x 100 m. One other specimen was taken in 1980, in similar prairie about 
12% miles north of Hays. It was in a disturbed area where numerous chalk fragments 
were scattered about the surface. 

On 26 May 1981 while collecting in southcentral Kansas, Ely and Rolfs found this 
species to be common in the sagebrush and sandsage prairie habitats at numerous local- 
ities in Comanche, Clark and Ford Counties. In sagebrush south of Clark State Fishing 
Lake (18 miles north of Ashland), it was exceeded in numbers only by Nathalis iole. A 
good series was collected, most of them as they fed at various species of yellow-flowered 
composites. The only specimen taken in Ellis County during 1981 was a single taken in 
the FHSU pasture by Guy Ernsting on 29 June. 

Vanessa annabella (Field).—Field (op. cit.) reported one from Scott County on 28 
October 1985. Ely collected single individuals over flower beds on the FHSU campus on 
6 October 1980 and 19 October 1981. Other immigrant species recorded at Hays during 
summer 1981 included Mestra amymone (2), Chlosyne lacinia (15+), Agraulis vanillae 
and Phoebis sennae (4). 

We wish to thank William H. Howe, who confirmed the identifications of the two 
state records and reviewed the manuscript. 


CHARLES A. ELY, MARVIN E. ROLFS AND THOMAS E. LABEDZ, Department of Biology, 
Forts Hays State University, Hays, Kansas 67601. 


Journal of the Lepidopterists’ Society 
37(3), 1983, 258 


ABERRANT GLAUCOPSYCHE LYGDAMUS COUPERI GRT. (LYCAENIDAE) 


On 11 June 1977, an aberrant female Glaucopsyche lygdamus couperi Grt. was col- 
lected at Mer Bleue Bog, near Ottawa, Ontario. This species is common in open areas 
throughout the Ottawa area wherever the hostplant, common vetch (Vicia spp.), grows. 
The specimen was collected in a field bordering the bog while it rested on the underside 
of a vetch leaf. 

Figs. 1, 2 show the ventral surfaces of a normal (Fig. 2), and the aberrant specimen 
(Fig. 1). In the aberrant specimen, the normal ventral postmedian row of spots is elon- 
gated, especially on the secondaries, where they form a series of roughly rectangular 
bars, as compared to the round or oval spots of the usual form. The specimen has been 
deposited in the Nova Scotia Museum collection, Halifax, Nova Scotia. I would like to 
thank Mary Primrose of Dalhousie University, Halifax, Nova Scotia, for photographing 
specimen. 


KENNETH NEIL, Department of Biological Sciences, Simon Fraser University, Bur- 
naby, British Columbia, CANADA V5A 1S6. 


£ 


Fics. 1 & 2. Females of Glaucopsyche lygdamus couperi, ventral surface: 1, aberrant 
specimen; 2, normal specimen. 


Journal of the Lepidopterists’ Society 
87(3), 1983, 259 


BOOK REVIEW 


LARGE WHITE BUTTERFLY: THE BIOLOGY, BIOCHEMISTRY AND PHYSIOLOGY OF PIERIS 
BRASSICAE (LINNAEUS). By John Feltwell. W. Junk Publishers, The Hague. Series Ento- 
mologica, vol. 18. 542 pp. 1981. $98.00. 


Large White Butterfly—it sounds like it is missing a “The” in front—is a very peculiar 
book. It attempts to be a complete, or nearly complete, literature survey of a very 
extensively studied animal, Pieris brassicae. It probably isn’t complete, but I failed to 
turn up any references in my own files that weren't in it—down to Akhmedov’s 1967 
paper in the proceedings of the Azerbaijanian Academy of Sciences on photoperiodic 
reactions of Tashkent stocks, or Fernando’s in Spolia Zeylandica on host plant selection. 
One can't get much more arcane than those. 

Why do a compilation like this? As Miriam Rothschild says in her foreword, the 
literature of everything is now very unwieldy. A few years ago I discovered that some 
experiments I had just published were foreshadowed precisely by Z. Lorkovié in work 
published in the scientific yearbook of the University of Zagreb in 1928. Missing that 
reference was fairly easy, but even keeping up with current journals in the library is a 
nightmare, and those able to do so are increasingly subscribing to computerized litera- 
ture-search services. Even that has its perils—if you try it with “Large White” as a search 
word, as I did, you will end up with dozens of papers on the culture of a breed of swine 
by that name, popular in Eastern Europe. Remarkably, Feltwell did this compilation 
without computerized assistance. Probably no one will ever do such a job that way again. 

Anyone working on any aspect of pierid biology will have recourse to this book for 
many years to come. Pieris brassicae is the most-studied pierid on most fronts—Colias 
eurytheme is probably next in line. The extent to which it is legitimate to extrapolate to 
other pierids, or even to other species placed in the old genus Pieris, is problematical; 
brassicae and its very close relatives form an odd, isolated pocket in the group, distin- 
guished by a very reduced chromosome number and other things. Still, it is all we ve got 
to compare most things to. 

The book is obviously a compilation. The information conveyed is telegraphic, frag- 
mentary, and often so out of context that no real picture of its significance emerges. Of 
course, much of the information conveyed is trivial, but unless one already knows the 
field one might be hard-pressed to tell what is important and what is not—in a way, 
Large White Butterfly is organized like an organic chemistry course, in which, to be 
safe, you learn everything. Information on aberrations and the like is dated and pretty 
meaningless when their etiology and developmental context are unexplored. They might 
be very interesting to developmental biologists, physiologists, or geneticists, but how is 
one to tell? 

This is not a coffee-table book. Despite its price, which works out to 18.3¢ a page, 
there arent even any color plates (But why should there be, in a book about a black- 
and-white “bug?’’). The book belongs in all major institutional reference libraries, where 
it will save graduate students, especially, a great deal of travail. If it fails to convey much 
impression of what the Large White really is, how it lives, and how it fits into its 
environment, it at least points the way to the original sources. The moral is simple: If 
you want character development, don't read the phone book like a novel. 


ARTHUR M. SHAPIRO, Department of Zoology, University of California, Davis, Cal- 
ifornia 95616. 


Journal of the Lepidopterists’ Society 
37(3), 1983, 260-261 


OBITUARY 


MURRAY OTTO GLENN (1893-1981) 


Murray Glenn (1977)—photo by G. L. Godfrey 


Microlepidopterology lost one of its most dedicated and productive, but also unsung, 
workers when Murray Glenn died in a hospital in Spring Valley, Illinois on 26 July 1981. 
He was not well known to many Lepidopterists, yet Dr. J. F. Gates Clarke (in litt.) 
characterized him as having “contributed more to our knowledge of midwestern micro- 
lepidoptera with particular reference to Illinois than any other person except, perhaps, 
the late Dr. Annette Braun.” 

Murray Otto Glenn was born in Magnolia, Illinois on 3 October 1893, the son of Isaac 
and Helen (Otto) Glenn. He married Lena F. Bell on 21 May 1925 in Normal, Illinois. 
She survives him together with three sons, Alan, Richard and Donald; a daughter, Mar- 
jorie (Mrs. George Haws); a sister, Mrs. Gladys Steer; 12 grandchildren and four great- 
grandchildren. A brother and a sister preceded him in death. 

Glenn studied agriculture at the University of Illinois for three years after graduating 
from high school in 1911 but withdrew before taking a degree to join the Army during 
WWI. He left the service as a lieutenant. As a young man he was hired by a railroad to 
play semi-professional baseball in Montana to entertain the settlers there. Returning to 
his native state, he farmed over 600 acres with his brother near Magnolia. After retiring 
from farming he moved to Henry, Illinois where he spent the rest of his life. 


VOLUME 87, NUMBER 3 261 


He began to collect insects, particularly Coleoptera, as a high school student, but his 
serious study of Lepidoptera began in 1931 when his interest was attracted to their 
immatures. In rearing these he discovered many life histories and foodplant associations. 
He collected the holotypes of 18 new species of microlepidoptera and paratypes of 10 
more, all described by other workers, and three of which were named in his honor. 
Several of these new taxa were described by Clarke, who said in further tribute to Glenn’s 
work (in litt.) that “his specimens were beautifully prepared thereby facilitating study. 
It was always a pleasure to work with his material.’’ Glenn published nothing on Lepi- 
doptera during his long career; his only publication was as junior author with Dr. M. W. 
Sanderson of a note on Coleoptera (1963, Coleopterists’ Bulletin 7:52). 

Glenn occasionally collected in other parts of the United States, but by far the greater 
portion of his field work was done in the immediate vicinity of his home. His efforts 
produced a virtually complete record of the microlepidoptera of that portion of Illinois, 
especially useful in monitoring environmental changes over a 46-year period. Although 
microlepidoptera were his specialty, he did not neglect the larger Lepidoptera, including 
butterflies. He professed a humorous disdain for the latter, but nevertheless turned up a 
number of Illinois rarities. 

Glenn continued his activities well into later life. He once told me that he had consid- 
ered giving up moth work when he reached 80, but when he did he “couldn't see that 
80 was any different than 79, so I am still doing what I enjoy most, and am collecting 
again this year.” In 1969 he donated his macrolepidoptera to the Illinois Natural History 
Survey (“Survey’) except for some common butterflies which he kept for show and 
eventual donation to a local school. In 1977 he turned over his identified micros to the 
Survey and his undetermined material to the United States National Museum of Natural 
History. The Survey regards the Glenn collection as the most important of the several 
Lepidoptera collections in its care. 

I became acquainted with Murray as a young man early in 1947, at about the same 
time that the Lepidopterists’ Society, which we both joined as charter members, was 
being formed. Before meeting him my outside contacts had been few, and through him 
I made new ones and broadened my horizons. We remained lifelong friends. He was 
generous, modest, humorous and always kind and helpful; a true “handmaiden of sci- 
ence,’ who will be missed by all who had the honor and pleasure of knowing him. 

I am very grateful to Dr. George L. Godfrey of the Survey for his help in the writing 
of this article. He published additional information on Glenn’s collection and activities, 
including a list of 17 of the taxa based on Glenn holotypes (1978, J. Lepid. Soc. 32:235) 
(the 18th name remains in manuscript at this writing). An article on Glenn’s life and 
work, some of which is excerpted herein by permission, was published by Godfrey and 
W. E. LaBerge (1977, Illinois Natural History Survey Reports No. 168). Family data 
were derived from Glenn’s obituary (replete with errors concerning his scientific work) 
in the Henry, Illinois News-Republican of 29 July 1981. The photograph was taken by 
Godfrey in 1977 at the time of the transfer of Glenn’s collection to the Survey. 


RODERICK R. IRWIN, 24 East 99th Place, Chicago, Illinois 60628 (Research Affiliate, 
Illinois Natural History Survey and Illinois State Museum). 


Journal of the Lepidopterists’ Society 
37(3), 1983, 262-264 


OBITUARY 


ERNST J. DORNFELD (1911-1983) 


Dr. Ernst J. Dornfeld (1911-1988), author of “The Butterflies of Oregon” (1980) (photo 
taken Nov. 1979 by John Neyhart). 


Ernst J. Dornfeld died on 30 May 1983 in Corvallis, Oregon at the age of 72. A 
professional cell biologist, he had been chairman of the Department of Zoology at Oregon 
State University for 24 years prior to his retirement in 1976. He was interested in but- 
terflies and skippers with a special interest in the fauna of Oregon, culminating in his 
book on the Butterflies of Oregon, published in 1980. 

Ernst was born in Milwaukee, Wisconsin on 6 April 1911, the son of Ernst Phillip 
Dornfeld and Gertrude Dornfeld. His father, a Lutheran minister, encouraged him when 
he showed interest in butterflies at an early age. He attended Concordia College and 
then Marquette University, receiving his B.S. degree from Marquette in 1933. He began 
his graduate work at the same institution, then moved on to the University of Wisconsin, 
at Madison, where he received his M.A. degree in 1935 and his Ph.D. in 1937. He spent 
the summer of 1935 at the University of Michigan, Ann Arbor. 


VOLUME 37, NUMBER 3 263 


His graduate research work was in cell biology and occupied most of his time during 
that period. He did serve as Assistant in Lower Zoology at the Milwaukee Public Museum 
from 1930 to 1932, while an undergraduate student at Marquette. Ernst often acknowl- 
edged the strong influence his germanic heritage exerted on his scientific work and on 
his interest in the liberal arts. It was his feeling that his broad interests were encouraged 
by his family and by the many cultural activities in Milwaukee, including the Milwaukee 
Public Museum. This institution continues to occupy a prominent position in the cultural 
life of Milwaukee. 

Ernst spent a year as an instructor in histology and embryology at the University of 
Oklahoma School of Medicine before coming to the then Oregon State College in 1938 
as an instructor of Zoology. He moved up through the academic ranks to Assistant 
Professor in 1942, Associate Professor in 1946, Professor in 1950 and Chairman of the 
Department of Zoology in 1952, a position he held until his retirement from Oregon 
State University in 1976 as Professor Emeritus of Zoology. He married Lorena Sue 
Ferguson in 1945. They had five children, Ernst, Susan, Ruth, Margaret and Carl. 

Ernst had a long and distinguished career as an academician and as a cell biologist. 
When he joined the department in 1938, there were six faculty members. When he 
retired in 1976, the department had grown to 17 faculty members. He was an unusually 
fine teacher, well organized, thorough, articulate and enthusiastic. During his years on 
the faculty, he taught eight different courses ranging from general zoology to the biology 
of the cell. He was well regarded by his students, receiving the Loyd Carter Award for 
“Inspirational Teaching” in 1947, the second such award to be given. 

Ernst was an accomplished research scientist besides a teacher. His research centered 
on cell biology, particularly cell division and differentiation. He published 47 scientific 
papers, including 9 on the Lepidoptera. It is of interest to note that his first and last 
papers were on butterflies. Ernst received the Society of Sigma Xi Research Award in 
1961, the Distinguished Professor Award from the Oregon State University Alumni As- 
sociation in 1978 and the Citation for Outstanding Scientific Achievement from the 
Oregon Academy of Science in 1981. While his earlier awards were based largely upon 
his work in cell biology, gradually his expertise with the Lepidoptera became better 
known and added to his stature as a scientist, culminating in the appearance of his book 
on Oregon butterflies. 

His interest in butterflies extended back to his childhood days and the encouragement 
he received from his family. He amassed a large collection of Wisconsin butterflies prior 
to coming to Oregon. Some of these specimens were contributed to the Milwaukee Public 
Museum at the time. His interest in Lepidoptera remained throughout his life but other 
research interests, particularly in cell biology, dominated his professional career for many 
years. His interest in butterflies was revived when one of his sons became interested in 
them in the early 1950's. Ernst began to study and collect the Oregon fauna with the 
same zeal and enthusiasm that he brought to his other work. He made numerous field 
trips into all parts of Oregon and gradually built up a magnificent collection of approx- 
imately 25,000 specimens, largely from Oregon. He compiled meticulous records of his 
own material and that of others against the day when he would have the time to complete 
his book on the fauna of Oregon. His retirement in 1976 provided him with that time 
he needed. He started to work on that book almost on the day he retired. It appeared in 
1980, just the way he wanted it to be. He was able to incorporate the summed knowledge 
of each species found in the state, based on his own work and that of others. There were 
many others, for Ernst established and maintained contact with a number of lepidop- 
terists, freely sharing his own knowledge and experience. His willingness to share and 
his enthusiasm was extended to a good many children in Corvallis, including the three 
sons of one of us (J.D.L.), who quickly learned that Ernst knew all there was to know 
about butterflies. Often, it was difficult to determine who was more excited over an 
interesting catch, Ernst or the youthful entomologist! 

Ernst delivered the annual Sigma Xi lecture in 1978 entitled “About Butterflies.”’ It 
was a beautifully prepared and illustrated lecture and was very well received. It was so 
well received, in fact, that shortly before he died, he told us that he had given that talk 
26 times since 1978. He was a member of the Lepidopterists’ Society for many years 


264 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


and served as co-host for the 1967 meeting held at Oregon State University. He was an 
active participant in the various lepidopterists’ workshops held around the northwest. His 
last active role was to host such a workshop in November of 1982 at Oregon State. 

The Dornfeld collection, including his library and records, has been bequeathed to the 
Systematic Entomology Laboratory (SEL), Department of Entomology, Oregon State 
University. The SEL contains approximately 2.4 million specimens, including the largest 
collection of butterflies in the Pacific Northwest. Ernst donated thousands of hours to the 
SEL. He worked over the entire collection of butterflies in his meticulous fashion, even 
leaving room throughout the collection for his own specimens. He was very active in 
encouraging others to donate their collections as well, resulting in thousands of additional 
specimens for the collection. He took his scientific responsibilities very seriously. 

While he was serious about his science, he was also infectiously enthusiastic and had 
a wonderful sense of humor. He was a perfectionist in everything he did but enjoyed 
nothing more than a joke upon himself. Both of us shared field and laboratory experiences 
with him for many years. We remember his warmth and friendship and especially his 
chuckle that often erupted into a loud laugh. Ernst had a love of life and learning and 
a real passion for butterflies. It was our privilege to have shared a part of his life. 

Our thanks to John Neyhart, Roseburg, Oregon, for the fine photograph of Ernst. 


Lepidoptera Publications by Ernst J. Dornfeld 


1931 Dornfeld, Ernst J. A night-flying butterfly and some unusual locality records (Lep- 
idoptera). Entomol. News 42:287. 

1960 Dornfeld, Ernst J. Mitoura johnsoni in Oregon and California. J. Lepid. Soc. 13: 
183. 

1962 Dornfeld, Ernst J. Butterflies of Oregon. Bull. Oregon Entomol. Soc. 8:50. 

1964 Dornfeld, Ernst J. Favorite collecting spots—The Ochoco Mountains. Bull. Oregon 
Entomol. Soc. 16:123-125. 

1967 Dornfeld, Ernst J. On the yellow forms of Coenonympha tullia (Satyridae) in 
Oregon. J. Lepid. Soc. 21:1-7. 

1970 Dornfeld, Ernst J. A field-captured scale-deficient mutant of Anthocaris sara. J. 
Res. Lepid. 9:25-28. 

1971 Dornfeld, Ernst J. & John Hinchliff. Check List of Oregon Bhoeaiocees with Coun- 
ty Records and Flight Periods. [2] + 7 + 5 pages, 1 map. Corvallis, Oregon: Dorn- 
feld and Hinchliff. 

1980 Dornfeld, Ernst J. The Butterflies of Oregon. Timber Press, Forest Grove, Oregon. 
XIV, 276 pp. 4 colored plates, 48 black-and-white plates, 192 maps. 

1983 Hammond, Paul C. & Ernst J. Dornfeld. A new subspecies of Speyeria egleis 
(Nymphalidae) from the pumice region of Central Oregon. J. Lepid. Soc. 37:115- 
120. 


JOHN HINCHLIFF, 2960 SW Bennington Drive, Portland, Oregon 97201 AND JOHN 


D. LATTIN, Systematic Entomology Laboratory, Department of Entomology, Oregon 
State University, Corvallis, Oregon 97331. 


Date of Issue (Vol. 87, No. 3): 27 April 1984 


= tions. 


EDITORIAL STAFF OF THE JOURNAL 
THOMAS D. EICHLIN, Editor 


% Insect Taxonomy Laboratory 
1220 N Street 
Sacramento, California 95814 U.S.A. 


MaGDA R. Papp, Editorial Assistant 


_ Douctas C. FERGUSON, Associate Editor THEODORE D. SARGENT, Associate Editor 


NOTICE TO CONTRIBUTORS 
_ Contributions to the Journal may deal with any aspect of the collection and study of 
Lepidoptera. Contributors should prepare manuscripts according to the following instruc- 


Abstract: A brief abstract should precede the text of all articles. 
Text: Manuscripts should be submitted in triplicate, and must be typewritten, en- 


| tirely double-spaced, employing wide margins, on one side only of white, 8% x 11 inch 


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_ plant or animal in the text should include the full scientific name, with authors of 


zoological names. Insect measurements should be given in metric units; times should be 
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_ italics are intended. References to footnotes should be numbered consecutively, and the 


footnotes typed on a separate sheet. 

Literature Cited: References in the text of articles should be given as, Sheppard 
(1959) or (Sheppard 1959, 1961a, 1961b) and all must be listed alphabetically under the 
heading LITERATURE CITED, in the following format: 


3 SHEPPARD, P. M. 1959. Natural selection and heredity. 2nd. ed. Hutchinson, London. 


209 pp. 
196la. Some contributions to population genetics resulting from the study of 
the Lepidoptera. Adv. Genet. 10: 165-216. 


In the case of general notes, references should be given in the text as, Sheppard (1961, 


co lh we 


ro 


ee SCC (iC 


—~ = 
So os 


q Adv. Genet. 10: 165-216) or (Sheppard 1961, Sym. R. Entomol. Soc. London 1: 23-30). 


Illustrations: All photographs and drawings should be mounted on stiff, white back- 
ing, arranged in the desired format, allowing (with particular regard to lettering) for 
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are not acceptable and should be reduced photographically to that size or smaller. The 
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PRINTED BY THE ALLEN PRESS, INC., LAWRENCE, KANSAS 66044 U.S.A. 


CONTENTS 


A NEw CLEARWING MOTH (SESIIDAE) FROM CENTRAL AMERICA: 
A STEM BORER IN MIMOSA PIGRA. Thomas D. Eichlin & 
Steven Passo i Us I 


VARIATION AND HOST SPECIFICITY IN THE YUCCA MOTH, TE- 
GETICULA YUCCASELLA (INCURVARIIDAE): A MORPHOMET- 
RIC APPROACH: Nancy Jo Miles 00000 2 
IMMATURE STAGES OF ANACAMPTODES HERSE (SCHAUS) (GEO- 
METRIDAE) ON SOYBEAN IN HONDURAS. Steven Passo ........ 
NEw Host RECORDS FOR OLETHREUTINAE (TORTRICIDAE). 
Richard L. Brown, J. F. Gates Clarke & Dale H. Habeck ... 
NEW WISCONSIN BUTTERFLY RECORDS. Roger M. Kuehn .......... 
PUPAL COLOR DIMORPHISM IN CALIFORNIA BATTUS PHILENOR 
(L.) (PAPILIONIDAE): MORTALITY FACTORS AND SELECTIVE 
ADVANTAGE. S. R. Sims ¢ A. M, Shapiro 2 a 
A NEW SUBSPECIES OF SPEYERIA ATLANTIS (EDWARDS) 
(NYMPHALIDAE) FROM THE GREAT BASIN OF NEVADA. 
George\T) Austin: 


THE LARVA OF SIDERIDIS MARYX (GUENEE) (NOCTUIDAE). 


Kenneth A. Neil SO ee BRUNA se le 


GENERAL NOTES 
Collection records of butterflies from Nyangezi, Zaire, Africa. William Gluck 
Notes on the genus Imelda (Riodininae). Curtis J. Callaghan _. 


New and unusual butterfly records from Kansas. Charles A. Ely, Marvin E. 
Rolfs <> Thomas En Dabedz) 20.000 UN No 


Aberrant Glaucopsyche lygdamus couperi Grt. (Lycaenidae). Kenneth Neil 
BOOK REVIEW) vo co UUse oy MS OO 


OBITUARIES 02. hae ay) Pa NUS UMeNP STUUR Suse e\ | 206, 260, 


207 4 Y 
PA 

224 
228 


236 


256) 
258 
259 


_ Volume 37 198s = Number 4 


haya | 


ISSN 0024-0966 


JOURNAL 


_ Lepipoprerists’ SociETy 


Published quarterly by THE LEPIDOPTERISTS’ SOCIETY 
 Publié par LA SOCIETE DES LEPIDOPTERISTES 


Herausgegeben von DER GESELLSCHAFT DER LEPIDOPTEROLOGEN 
Publicado por LA SOCIEDAD DE LOS LEPIDOPTERISTAS 


4 
‘ DEDICATED TO THE MEMORY OF 
K Lucien Harris, Jr. 


1899-1983 


8 May 1984 


THE LEPIDOPTERISTS’ SOCIETY 


EXECUTIVE COUNCIL 


LEE D. MILLER, President CHARLES V. COVELL, JR., 
KAROLIS BAGDONAS, Vice President Immediate Past President 
MIGUEL R. GOMEZ BUSTILLO, Vice President JULIAN P. DONAHUE, Secretary 
J. DONALD LAFONTAINE, Vice President RONALD LEUSCHNER, Treasurer 


Members at large: 


K. S. BROWN, JR. F. S. CHEW J. M. BuRNS 
E. D. CASHATT G. J. HARJES F. W. PRESTON 
T. C. EMMEL E. H. METZLER N. E. STAMP 


The object of the Lepidopterists’ Society, which was formed in May, 1947 and for- 
mally constituted in December, 1950, is “to promote the science of lepidopterology in — 
all its branches, . . . . to issue a periodical and other publications on Lepidoptera, to facil- 
itate the exchange of specimens and ideas by both the professional worker and the 
amateur in the field; to secure cooperation in all measures’ directed towards these aims. 

Membership in the Society is open to all persons interested in the study of Lepi- 
doptera. All members receive the Journal and the News of the Lepidopterists’ Society. 
Institutions may subscribe to the Journal but may not become members. Prospective 
members should send to the Treasurer full dues for the current year, together with their 
full name, address, and special lepidopterological interests. In alternate years a list of 
members of the Society is issued, with addresses and special interests. There are four 
numbers in each volume of the Journal, scheduled for February, May, August and 
November, and six numbers of the News each year. 


Active members—annual dues $18.00 
Student members—annual dues $12.00 
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Send remittances, payable to The Lepidopterists’ Society, and address changes to: 
Ronald Leuschner, 1900 John St., Manhattan Beach, California 90266 U.S.A. 


Back issues of the Journal of the Lepidopterists’ Society, the Commemorative Vol- 
ume, and recent issues of the NEWS are available from the Treasurer. The Commem- 
orative Volume, is $6; for back issues, see the NEWS for prices or inquire to Treasurer. 


Order: Mail to Ronald Leuschner, 1900 John St., Manhattan Beach, California 90266 
U.S.A. 


Journal of the Lepidopterists’ Society (ISSN 0024-0966) is published quarterly by the 
Lepidopterists’ Society, a non-profit, scientific organization. The known office of publi- 
cation is 1041 New Hampshire St., Lawrence, Kansas 66044. Second class postage paid 
at Lawrence, Kansas, U.S.A. 66044. 


Cover illustration: Adult of the squash vine borer, Melittia cucurbitae (Harris) (Sesi- 
idae), which occurs in the eastern half of the United States and along the Gulf Coast into 
Vera Cruz, Mexico. The larvae are destructive borers in the vines of various cultivars of 
Cucurbita spp. (squash, pumpkins and gourds). Original drawing by Dr. Charles S. Papp, 
Sierra Graphics & Typography, 1722 J Street #19, Sacramento, CA 95814, USA. 


JoURNAL OF 
Tue LEPIDOPTERISTS’ SOCIETY 


Volume 37 1983 Number 4 


Journal of the Lepidopterists’ Society 


 87(4), 1983, 265-268 


CYPRIPEDIUM FLOWERS ENTRAP ADULT THYMELICUS 
(LEPIDOPTERA: HESPERIIDAE) IN 
NORTHERN MICHIGAN 


EDWARD M. BARROWS 


Department of Biology, Georgetown University, 
Washington, D.C. 20057 


ABSTRACT. Adults of the introduced skipper Thymelicus lineola were attracted to 
the nectarless flowers of the native orchids Cypripedium reginae and C. calceolus. No 
doubt in search of food, they crawled into orchid labella. Up to 24 skippers and other — 
insects became entrapped and died in a single labellum. 


The European skipper, Thymelicus lineola (Ochsenheimer), was first 
discovered in North America in London, Ontario, Canada, in about 
1910 (Saunders, 1916; Klots, 1958). Since this time it has spread to 
British Columbia, New Brunswick, Manitoba, Connecticut, New Jer- 
sey, New York, Pennsylvania, Ohio, Michigan, Maryland, and Virginia 
(Burns, 1966, pers. comm.; Preston & Westwood, 1981) and has become 
a pest of hay fields in Canada (McNeil & Duchsne, 1975; McNeil et 
al., 1976). Adults seek nectar from many species of flowers including 
the lady’s-slipper orchids Cypripedium reginae Walter and C. calceo- 
lus L. This report increases knowledge about the peculiar entrapment 
of adult T. lineola by the labella of these orchids in northern Michigan. 
Arthur (1962) previously reported this phenomenon regarding this but- 
terfly and C. reginae in Ontario, and Catling (1974) reported it in both 
Ontario and southern Michigan. 

Observations were made in summer, 1977, in a marshy and swampy 
area on the property of the University of Michigan Biological Station, 
Cheboygan County, Michigan, when the skipper was abundant. Orchid 
labella that were partially or wholly dried after anthesis were removed 
from pedicels for inspection of their contents. Thymelicus lineola was 
uncommon in the study area in the summer of 1978 when I attempted 
to continue this study. 


266 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Fic. 1. A Thymelicus lineola probing the labellar orifice of a nectarless flower of 
Cypripedium calceolus. 


Thymelicus lineola frequently alighted on orchid flowers, extended 
their proboscides into labellar openings and crawled into the labella 
(Fig. 1). They apparently obtained no food from these nectarless flow- 
ers (Stoutamire, 1967; pers. obs.). Cypripedium labella are adaptations 
for bee, not butterfly, pollination; bees may become temporarily en- 
trapped in them and effect pollination by depositing pollinia as they 
escape through one of two small orifices at the labella bases (Stouta- 
mire, 1967; Catling, 1974). 

On 26 June, 219 T. lineola were found in 42 C. reginae flowers. 
Each flower contained from 0 to 11 males (x = 5) and from 0 to 4 
females (x = 0.5). On 4 July, 42 flowers contained 427 dead T. lineola, 


~ VOLUME 37, NUMBER 4 267 


with from 0 to 15 males (x = 7.4) and from 0 to 12 females (x = 2.7) 
per flower. The flowers contained significantly more males (90.41%) 
on 26 June than on 4 July (73.0%) (P = 0.035, test for equality of two 
percentages). Up to 24 T. lineola were found in a single labellum. 
Catling (1974) reported up to five of them per labellum. In late June, 
I discovered a female acridid grasshopper, Melanoplus islandicus 
Blatchley, eating dried T. lineola that she obtained through a hole in 
a dried labella. These flowers also contained salticid and thomisid spi- 
ders; entomobryid springtails; perlodid stoneflies; reduviid and mirid 
bugs; derodontid, elaterid, and lathridiid beetles; anisopodine, chloro- 
pid, phorid, and syrphid flies; geometrid moths; and andrenine bees. 
Most of these arthropods were dead. 

On 26 June and 4 August, a total of eight of 11 inspected C. calceolus 
flowers contained insects. From one to three males of T. lineola were 
in four of the flowers and one female T. lineola was in one flower. 
These flowers also contained a thomisid spider, a culicid fly, geometrid 
moths, lathridiid beetles, and halictine, andrenine and megachilid bees. 

The native skippers, Polites themistocles Latreille and P. coras (Cra- 
mer), also enter Cypripedium labella; however, they were not found 
entrapped in them (Guignard, 1886; pers. obs.). In comparison to T. 
lineola, these native butterflies may be able to escape from the labella 
due to their greater strength, behavioral flexibility, or both. Catling 
(1974) suggested that Ewphyes and Ancyloxipha skippers flying in a 
bog with C. reginae in Ontario may not be trapped due to their having 
feeding habits different from T. lineola. Further, he made the plau- 
sible hypothesis that fatal entrapment of T. lineola is an ‘“‘accident”’ 
due to their encountering a North American orchid with characteristics 
of C. reginae. Finally, he surmised that pollination of this orchid may 
be reduced by entrapment of T. lineola because they could obstruct 
pollinator movements. These and other interesting hypotheses regard- 
ing this skipper and these orchids remain to be tested. 


ACKNOWLEDGMENTS 


Edward G. Voss (University of Michigan), John M. Burns (Smithsonian Institution), 
and an anonymous reviewer made helpful suggestions regarding this article. Ashley B. 
Gurney (USDA, Washington, D.C.) identified the Melanoplus. 


LITERATURE CITED 


ArtTHur, A. P. 1962. Adults of the European skipper, Thymelicus lineola (Ocsh.) 
(Lepidoptera: Hesperiidae) trapped in flowers of the showy lady’s slipper orchid. 
Proc. Entomol. Soc. Ontario 92:190-191. 

BURNS, J. M. 1966. Expanding distribution and evolutionary potential of Thymelicus 
lineola (Lepidoptera: Hesperiidae), an introduced skipper, with special reference to 
its appearance in British Columbia. Can. Entomol. 98:859-866. 


268 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


CATLING, P. M. 1974. A butterfly-trapping orchid. Newsletter Mich. Entomol. Soc. 19: 
I, ®s 

GUIGNARD, J. A. 1886. Insects and orchids. Ann. Rep. Entomol. Soc. Ontario 16: 
39-48. 

KiLots, A. B. 1958. A Field Guide to the Butterflies of North America, East of the 
Great Plains. Houghton Mifflin, Boston. 349 pp. 

MCNEIL, J. N. & R. DUCHSNE. 1975. Known distribution of the European skipper, 
Thymelicus lineola (Lepidoptera: Hesperiidae), in Quebec. Can. Entomol. 107:1221- 
1223) 

MCNEIL, J. N., S. A. TURNBULL & C. R. Harris. 1976. Laboratory studies on the 
contact toxicity of some insecticides to the European skipper, Thymelicus lineola 
(Lepidoptera: Hesperiidae). Can. Entomol. 108:319-320. 

PRESTON, W. B. & A. R. WESTWOOD. 1981. The European skipper, Thymelicus lineola 
(Lepidoptera: Hesperiidae), in Manitoba and northwestern Ontario. Can. Entomol. 
113:1129-1130. 

RAWSON, G. W. 1931. The addition of a new skipper, Adopaea lineola (Ocsh.), to the 
list of U. S. Lepidoptera. J. New York Entomol. Soc. 39:503-506. 

SAUNDERS, W. E. 1916. European butterfly found at London, Ont. Ottawa Nat. 30: 
116. 

STOUTAMIRE, W. P. 1967. Floral biology of the lady’s-slippers (Orchidaceae: Cypri- 
pedium). Mich. Bot. 6:159-175. 


Journal of the Lepidopterists’ Society 
37(4), 1983, 269-274 


SIGNIFICANCE OF VISITS BY HACKBERRY BUTTERFLIES 
(NYMPHALIDAE: ASTEROCAMPA) TO FLOWERS 


RAYMOND W. NECK 


Texas Parks & Wildlife Department, 4200 Smith School Road, 
Austin, Texas 78744 


ABSTRACT. Behavioral studies were conducted on hackberry butterflies (Astero- 
campa spp.) in central Texas as to their visitations to certain flowers. The butterflies were 
observed most abundantly on snakewood flowers (Colubrina texensis). A higher per- 
centage of females were counted during sampling. It is speculated that Asterocampa 
spp., especially females, visit flowers which serve as a nitrogen source and do not require 
the carbohydrate-rich nectar sources of most other flowers. 


Visits by rhopaloceran species to angiospermous flowers are well 
known even to laymen. However, as any knowledgeable lepidopterist 
is aware, some species are rarely or never observed at flowers. One 
group of butterflies generally believed not to visit flowers are the hack- 
berry butterflies of the nymphalid genus Asterocampa. Howe (1975: 
118) states that “they do not visit flowers but feed on decaying mate- 
rial—rotting fruit, fermenting tree sap, animal excrement and car- 
casses.” No record of Asterocampa visiting flowers was known to Shields 
(1972) who reviewed butterfly flower visitation records. The most ex- 
tensive published account of the life history account of any Asterocam- 
pa appears to be that of A. celtis (Langlois & Langlois, 1964). Adults 
were observed feeding at wet mud and fruit (mulberries and cherries). 
Kimball (1965) recounted observations of Asterocampa spp. feeding at 
rotton persimmons and oozing hackberry trees. A. clyton and A. celtis 
have been observed feeding at pig carrion in a state of “advanced 
decay” (Payne & King, 1969). Visits to waterholes in Arizona by A. 
celtis were observed by Bauer (1953). In Arizona, A. leila has been 
observed feeding at coyote feces which contained much Opuntia fruit; 
they were never observed at flowers or mud puddles (Austin, 1977). In 
the eastern United States, Shapiro (1966) reported that A. celtis was 
“never seen ... on flowers’ while A. clyton were “occasionally seen 
clustered on over-ripe fruit.” The only reference to Asterocampa on 
flowers that I have found is Scott and Scott (1980), who report that A. 
celtis in Colorado “rarely feed on flowers (Jamesia, etc.) but often feed 
on sap especially of willows and occasionally feed on mud or on Rubus 
berries.’ Kimball (pers. comm.), however, says he has observed Aster- 
ocampa spp. visiting a variety of flowers on several occasions. The 
purpose of this communication is to document significant visitations 
by Asterocampa spp. to angiospermous flowers and to elucidate envi- 
ronmental factors which resulted in this activity. 


270 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


OBSERVATIONS 


All observations discussed below, unless otherwise so indicated, were 
made in McKinney Falls State Park, Travis Co., Texas, just southeast 
of Austin. The specific area was an upland flat above the left bank of 
Onion Creek. Shallow soil covers the underlying Pflugerville Limestone 
(Upper Cretaceous: Gulf Series) except for isolated areas and the scarp 
where bedrock is exposed. Three vegetational associations are present 
at the site. The woodland consists of cedar elm, Ulmus crassifolia 
Nutt., with an occasional plateau live oak, Quercus fusiformis Small. 
A meadowlike open area is dominated by small bur-clover, Medicago 
minima (L.) L. and yellow stonecrop, Sedum nuttalianum Rof. A 
thicket association occurs at certain woodland edges and the scarp of 
the upland flat. Dominant shrubs of this thicket are snakewood, Col- 
ubrina texensis (T. & G.) Gray; Texas persimmon, Diospyros texana 
Scheele; spring herald, Forestierra pubescens Nutt.; and pink cat-claw, 
Mimosa borealis Gray. The only hackberry occurring in this imme- 
diate area are seedling, sapling and small shrubby netleaf hackberry, 
Celtis reticulata Torr. Approximately 50 m from the study site, tree- 
size (to 8 m) individuals of both C. reticulata and Texas sugarberry, 
Celtis laevigata Willd. are present. | 

Observations on the study site were initiated on 7 March 1977 in 
connection with studies on other rhopalocera. An occasional Astero- 
campa was observed as early as 19 April when one fresh specimen of 
A. antonia (Edwards) was observed. Open flowers of snakewood were 
initially observed on 18 April. Moderate numbers of this species and 
A. texana (Skimmer) were observed feeding at flowers of Colubrina 
(Rhamnaceae) as early as 25 April (see Table 1). Flowers of snakewood 
are rather inconspicuous, being about 5 mm in diameter. The reduced 
petals form a yellowish five-pointed star surrounding a central green 
disc. Subsequently, for over two weeks adults of both species were 
observed at snakewood flowers. Last observed Asterocampa feeding at 
snakewood flowers was 10 May, although open flowers persisted through 
at least 19 May at this site. Apparent age of the butterflies was quite 
variable. All of the A. texana were fresh in appearance, although an 
age of one or two weeks is not unlikely. Adults of A. antonia varied 
from fresh to worn, very tattered adults. These tattered A. antonia 
were predominantly males. 

Detailed behavioral observations of these butterflies were recomend 
A butterfly at a snakewood flower was always observed to be moving 
its proboscis over the surface of the bow] of the flower. Nectar was not 
the object of this behavior. Unidentified substances, possibly including 
rich amounts of various amino acids, were obtained from the green 


VOLUME 37, NUMBER 4 Pat | 


TABLE 1. Asterocampa observed at snakewood (Colubrina texensis) flowers in 1977. 


A. antonia A. texana 
fo) Q fc) g 
25 April — 3:0* — 0:1 
26 April 3:1 5:0 — Mel 
29 April 0:1 0:3 — 2:0 
6 May — 1:0 1:0 1:0 
10 May — — — 1:0 
Total SEY 9:3 1:0 Sal 


2 total—14:4; 6 total—4:2; A. a.—12:5; A. t.—6:1. 
* At flower : not at flower. Butterflies counted as “at flower” only if proboscis observed extended to disk of flower. 


central disk. That significant amounts of amino acids may be available 
from snakewood flowers is indicated by the presence of carrion and 
dung flies at these flowers. Flowers attracting such flies must utilize 
high-level amino acid solutions in order to lure them away from their 
normal food sources which are naturally high in nitrogen (Baker & 
Baker, 1973a, b, 1975). All butterflies listed in Table 1 were observed 
to perform such behavior. During such activities neither feet nor an- 
tennae of Asterocampa touched the flower. Detection of the flower is 
apparently made visually with verification involving the highly flexible 
tip of the proboscis. Occasionally, the antennae are flexed up and down 
in unison, but no physical contact of the antennae was made with the 
flower. Possibly olfactory receptors in the antennae are receptive to 
chemicals emanating from the flowers. Movement from flower to flow- 
er was accomplished by walking along branches. 

As only the proboscis of Asterocampa comes into contact with the 
flower, these butterflies are unlikely to be effective pollinators. From 
this standpoint, Asterocampa spp. can be considered to be “‘cheaters’’ 
or nutrient-thieving flower visitors (Heinrich & Raven, 1972), because 
they do not participate in pollen transport. Legitimate flower visitors 
(actual pollinators) included at least two paper wasps (Polistes annu- 
laris and P. apacheanus), 2 tachinid fly species, 1 ichneumon wasp, 1 
muscid fly, 2 syrphid fly species, 1 conopid fly, honey bees (Apis mel- 
lifera L.) and 1 blow fly. 

A number of other rhopaloceran species visited snakewood flowers, 
while several butterflies present in the area were never observed vis- 
iting these flowers (Table 2). Examination of Table 2 reveals a definite 
dichotomy between two classes of butterflies—those species which visit 
snakewood flowers and those that don’t. That this is an ecologically 
significant dichotomy is remarkably demonstrated by the behavior of 
pollinating agents of snakewood and of balsam gourd, Ibervillea lind- 
heimeri (Gray) Greene, a vine growing upon snakewood. Both plants 


104 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 2. Rhopalocera observed during survey. 


Species observed at snakewood flowers Species at other flowers 
Atlides halesus estesi Clench Erynnis funeralis (Scudder & Burgess) 
Panthiades m-album (Bdv. & Lec.) Battus philenor philenor (Linnaeus) 
Strymon melinus franki Field Phoebis sennae marcellina (Cramer) 
Libytheana bachmanii larvata (Strecker) Abaeis nicippe (Cramer) 
Asterocampa antonia (W. H. Edwards) Nathalis iole Boisduval 
Asterocampa texana (Skinner) Chlosyne lacinia adjutrix Scudder 
Vanessa atalanta rubria (Fruhstorfer) Agraulis vanillae incarnata (Riley) 


Danaus plexippus plexippus (Linnaeus) 
Danaus gilippus strigosus (Bates) 


were blooming simultaneously, but pollinators of the two plants formed 
two mutually exclusive groups. Asterocampa and other visitors to 
snakewood flowers totally ignored balsam gourd flowers, while visitors 
to balsam gourd flowers (e.g., the butterfly, Abaeis nicippe Cramer, 
and a bumblebee, Bombus sp.) totally ignored snakewood flowers. 
Flowers of the two species were no more than five centimeters from 
each other. 

Although Asterocampa were found abundantly on snakewood flow- 
ers, a limited number of other food sources were noted. On 29 April 
one female A. antonia was observed on pencil cactus, Opuntia lep- 
tocaulis DC.; the proboscis of this individual was actively probing at 
the internodal joints of this plant. On 10 May a female A. texana alit 
on my arm, extended its proboscis and probed along the skin surface. 
On this same day one male A. texana and one female A. antonia were 
observed visiting flowers of Canada garlic, Allium canadense L. var. 
canadense. On 18 May one male A. antonia was observed feeding at 
flowers of this same plant. On this same day one female A. texana was 
observed feeding at mud. On 20 May one male and one female were 
observed feeding at flowers of Acacia angutissima (Mill.) O. Ktze. var. 
hirta (Nutt.) B. L. Robinson. 

One female A. texana was observed feeding on rotting fruit of pur- 
ple leaf plum (Prunus cerasifera Ehrh. var. pissardii Koehne) in a 
residential yard in Austin. This butterfly appeared to be using its an- 
tennae to help locate fruit. The antennae were flexed up and down on 
the surface of the fruit in unison and were even whirled in complete 
circles in front of the body. However, widespread probing at the fruit 
with the proboscis was also observed. 

Most blossoms present during those observations were ignored by 
Asterocampa, however. These included Medicago minima (L.) L., 
Sedum nuttalianum Rof., Lesquerella recurvata (Gray) Wats., Gail- 
lardia pulchella Foug., Tradescantia ohioensis Rof., Phacelia congesta 


VOLUME 387, NUMBER 4 DIS) 


Hook., Cooperia drummondii Herb., Torilus nodosa (L.) Goert. and 
Zexmenia hispida (H.B.K.) Gray. Of particular interest were observa- 
tions of two female A. texana which were observed landing on flow- 
ering inflorescences of P. congesta after they were “‘frightened”’ away 
from snakewood flowers (one individual disturbed by Polistes apa- 
cheanus, one by author). Each butterfly began investigating the P. 
congesta inflorescence with its proboscis; this behavior did not last 
more than two or three seconds as the butterflies appeared to be almost 
repelled by some characteristic of these flowers. The butterflies flew 
off to a branch of snakewood following this behavior. 


DISCUSSION AND CONCLUSION 


Of vital importance to the elucidation of the significance of flower 
visitation by Asterocampa is the preponderance of females in the sam- 
ples observed. Of twenty-four butterflies actually verified to be feeding 
at snakewood flowers, eighteen or 75.0% were females. Normally, sam- 
ples of butterflies from flowers yield a preponderance of males. How- 
ever, these species are obtaining mostly carbohydrate from their nectar 
sources, although nectar of butterfly flowers are often fairly rich in 
amino acids (Baker & Baker, 1973b). Adult protein requirements are 
relatively low in these species, because nearly-sufficient amounts of 
nitrogen are obtained during its larval development. 

Some butterfly larvae, however, are probably not able to store suf- 
ficient nitrogen for reproductive efforts because of difficulty in extract- 
ing nitrogen from foodplant material. The phytochemical defense of 
many tree species involves the production of “quantitative” poisonous 
substances, e.g., tannins, resins and silicates (Feeny, 1976). Owen (1959) 
observed that a British satyrid butterfly, Pararge algeria (L.) rarely 
visits flowers; whereas, a congeneric species, P. megera (L.), frequently 
visits flowers. This was explained by habitat selection, the former species 
confined to woodlands and the latter to open areas, presumably because 
of habitat restriction of larval foodplant. Such an explanation may well 
be valid in the above case but does not hold for Asterocampa. While 
the Celtis utilized as larval foodplants occur in woodlands, these wood- 
lands are open associations and in no way form closed canopies except 
under certain conditions; carbohydrate-rich nectar sources are nearly 
always nearby. Lack of visits by Asterocampa to showy, “typical but- 
terfly” flowers (see list in text above), are caused by a lack of certain 
nutrients required by adult Asterocampa, especially females. 

Quite possibly, Asterocampa, because it feeds as a larva on an “ap- 
parent” foodplant (see Feeny, 1976), suffers from a nitrogen-deficient 
larval diet. As a result, the imago would have to acquire sufficient 
nitrogen in order to reproduce. As the female contributes more pro- 


274 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


toplasm to the eggs which initiate the next generation, nitrogen re- 
quirements for adult female Asterocampa would be expected to be 
higher than requirements for male Asterocampa. As females would 
then spend more time obtaining nitrogen, one would expect to find 
more females than males in a given time period at a nitrogen source, 
e.g., flowers of Colubrina texensis or snakewood. However, certain 
differences in adult energy budgets of male and female butterflies are 
known (Adler, 1982). 


LITERATURE CITED 


ADLER, P. H. 1982. Why do male butterflies visit mud puddles? Can. J. Zool. 60:322- 
325. 

AUSTIN, G. T. 1977. Notes on the behavior of Asterocampa leila (Nymphalidae) in 
southern Arizona. J. Lepid. Soc. 31:111-118. 

BAKER, H. G. & I. BAKER. 1973a. Amino acids in nectar and their evolutionary signif- 
icance. Nature 241:543-545. 

1973b. Some anthological aspects of the evolution of nectar-producing flowers, 

particularly amino acid production in nectar. Pp. 2438-264, in V. H. Heywood (ed.). 

Taxonomy and Ecology. Systematics Association (London), Spec. Vol. 5. 870 pp. 

1975. Studies of nectar-constitution and pollinator-plant coevolution. Pp. 100- 
140, in L. E. Gilbert and P. H. Raven (eds.). Coevolution of Animals and Plants. U. 
Texas Press, Austin. 

BAUER, D. L. 1953. Butterflies at water holes in central Arizona. Lepid. News 7:146- 
147. 

FEENY, P. 1976. Plant apparency and chemical defense. In J. W. Wallace and R. L. 
Mansell (eds.). Biochemical Interaction Between Plants and Insects. Recent Adv. 
Phytochemistry 10:1-40. 7 

HEINRICH, B. & P. H. RAVEN. 1972. Energetics and pollination ecology. Science 176: 
597-602. 

Howe, W.H. 1975. The Butterflies of North America. Doubleday, Garden City, New 
York. 

KIMBALL, C. P. 1965. Lepidoptera of Florida. Arthropods of Florida and Neighboring 
Land Areas, Vol. 1:363 pp. Fla. Dept. of Agr., Gainesville. 

LANGLOIS, T. H. & M. H. LANGLOIs. 1964. Notes on the life-history of the hackberry 
butterfly, Asterocampa celtis (Bdvl. & Lec.) on South Bass Island, Lake Erie. Ohio 
J. Sci. 64:1-11. 

OwEN, D. F. 1959. Ecological segregation in butterflies in Britain. Entomol. Gaz. 10: 
27-38. 

PAYNE, J. A. & E. W. KING. 1969. Lepidoptera associated with pig carrion. J. Lepid. 
Soc. 23:191-195. 

ScoTT, J. A. & G. R. Scott. 1980. Ecology and distribution of the butterflies of southern 
central Colorado. J. Res. Lepid. 17:73-128. 

SHAPIRO, A. M. 1966. Butterflies of the Delaware Valley. Spec. Publ. Amer. Entomol. 
SOC: e/9ipp: 

SHIELDS, O. 1972. Flower visitation records for butterflies (Lepidoptera). Pan-Pac. 
Entomol. 48:189-203. 


Journal of the Lepidopterists’ Society 
87(4), 1988, 275-280 


A TWELVE YEAR COUNT OF THREE CALIFORNIA 
BUTTERFLIES 


LESLIE V. SMITH 
7627 Sycamore Drive, Citrus Heights, California 95610 


ABSTRACT. A weekly population count of three northern California butterflies, 
Danaus plexippus, Nymphalis antiopa and Papilio rutulus, over twelve years was con- 
ducted in a lowland California suburban yard. The data support Shapiro’s theory that 
N. antiopa goes into hibernation locally about mid-June. They do not support the oc- 
casionally heard statement that the numbers of butterflies are declining; no systematic 
trend is evident, but year-to-year fluctuations are pronounced. A hypothesis that P. 
rutulus thrives on extremes of rainfall is proposed. 


Most Lepidopterists cannot afford the time to do long-term butterfly 
counts. This is a twelve year count of the northern California butter- 
flies, Danaus plexippus (Linnaeus, Danidae), Nymphalis antiopa (Lin- 
naeus, Nymphalidae), and Papilio rutulus (Lucas, Papilionidae), made 
in a suburban Citrus Heights, Sacramento County, yard. The dominant 
vegetation was Quercus wislizenii, Juglans hindsii, Fraxinus velutina, 
Catalpa speciosa, and a variety of unknown grasses. Flowers that at- 
tract butterflies are Vinca major, Verbena peruviana, Phlox spp., and 
Rhododendron spp. There are many other flowers that were rarely if 
ever visited. 

Citrus Heights lies in a Mediterranean climatic regime, with very 
high year-to-year variability in both seasonal precipitation and its tim- 
ing (Figgins, 1979). The study period (1970-1982) embraces the most 
extreme and variable northern California weather in the 20th century, 
and to the extent that weather influences butterfly population levels, 
which is at least a debatable point (Shapiro, 1979), the counts reported 
should reflect the range of variation of which these species are capable. 

The field of view was generally 100 m to the north, east, and south 
with obstructions of shrubs and trees 10 m to the northeast and south- 
east. Observations to the west were sporadic to 20 m. The sample was 
based on a two hour watch period starting between 1100 h and 1300 
h every day except for cold or wet periods. Of course there were some 
days missed or observation periods of less than two hours. The elevation 
was 50 m, and the count was by unaided eye. 

The numbers of butterflies actually seen in such a count could be 
affected by a very large number of factors, some of which relate to 
actual population levels, while others may not. Examples of the latter 
are mowing, pruning, and planting practices on adjacent properties, 
which could affect the spatial distribution of host plants and nectar 
sources, and thus individual dispersal as well. Since the individual but- 
terflies were not marked, the counts can at best be regarded as an 


276 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 1. Danaus plexippus two hour daily per week count for twelve years at an 
observation point in Citrus Heights, California, U.S.A. 


Date 1970 1971 1973 1974 1975 1976 1977 1978 1979 1980 1981 1982 


II /26-IlI/4 0 
I/5-11 
III/12-18 

III /19-25 

III /26-IV/1 
IV /2-8 
IV/9-15 

IV /16-22 

IV /23-29 
IV/30-V/6 
V/7-13 
V/14-20 

V /21-27 

V /28-V1/3 
VI/4-10 

WAY I 
VI/18-24 
V1/25-VII/1 
VII/2-8 
VII/9-15 
VII/16-22 
VII/23-29 
VII/30-VIII/5 
VIII/6-12 
VIII/13-19 
VIII /20-26 
VIII /27-1X/2 
IX /3-9 

IX /10-16 
1.6/2) One ee 
1X /24-30 Ae oe 
xo ae 
X/8-14 
X/15-21 

X /22-28 
X/29-X1/4 
XI/5-11 
XI/12-18 


Total 100 


SHI Ori) OOP OoooeoeFWwwoeeS 
WOWONWANDRrROONNOGONTGWCOrFrAOCCSO 


1 
SGSOOrWWERFOWANWUODDHRARORFOGOOGOOOCOOCOOOFONZDOOCCOCC SO 


_ 


— 
SGOODOONOWKFAFrPNNANTBRWNNFOORrFRFWOORFrHFY BNHONH WwW 
— 
aN 


— 
SOCOOCRrFWWWAONNHTAOAArFrSABHORrRNNFrOCOr Cc COOoOWwWooOrce 


_ 


See 
5 GD CGI O NINOS Lom OOPOMMOoOoL my. | | | 
= 


—a— 
WHDOUONOAOWTANOUNGDODOODOONKFOONrFrOCOOFOrFrOCOOCOC oO 
a 


= 
SOR FKOWFrOPNAOArPAHOrrFOOOrFOOrFONNRYHYNABFOOrFrOO 


| o | orf 
ODOOODOOORFAWNHBKKBBKOIDOODNONNAWDWANrFrOOCOArFKUCOOCO 


_ 
ee) 

SOOO OOWOKBNFRRrFHWHAIOWONWNNORrFNNBKFONDOOWWNWOFH: 

SODOCDOWANDONAWANMNIONRrFOWBODOOFRrWANODOCOrFrRFGOCOCOOCOOCO SO 


oooorrs 
SOOO eC eS 
ONOWkOWKOUN WOOD 


~] 
S 
~] 
I 
(2) 
oo 
(ep) 
Co 


147 


> 
=) 
ice) 
(eX) 
(ee) 
aay 
ie) 
1-2) 
~] 
oO 
~] 
i) 


index of local abundance and certainly not as literal population esti- 
mates. Nonetheless, they are definitely useful. Throughout the twelve 
years there were no major disturbances in the surrounding vegetation 
or land-use patterns, and the count methods, however idiosyncratic, 
were thoroughly consistent. The data are thus indicative of local abun- 
dance, and similar long runs of such data are rare in the butterfly 
literature. They are largely consistent with trends observed in the same 
geographic area, using more extensive sampling methods by A. M. 
Shapiro (pers. comm.) between 1972 and 1982. 


VOLUME 87, NUMBER 4 2a 


TABLE 2. Nymphalis antiopa two hour daily count per week for twelve years at an 
observation point in Citrus Heights, California, U.S.A. 


Date 1970 1971 1973 1974 1975 1976 1977 1978 1979 1980 1981 1982 
1/16-21 a a le Oe ee LO. 0 
1/22-28 a ee ee) O; = HORE TO 
1/29-11/4 ary Oi — A lah teh Op. On eniOe) peerllg cee 
II/5-11 I iti Nt AO lla 210 | caluaey S on Ol agsO 
1112-18 be hs Ol ee OY, OM On Ore a ON el 
II/19-25 ee Oe Oe ee OP Or Oy OR CO. Aes 0 
II /26-Il1/4 ME OES Ong 0) ON CA SOs OS SOe Alo Leal 
Ill /5-11 EF 0yy (Os) Om, (ORY, Qu Os Sie, O. aeGrn gel 
III/12-18 ee) Ome Op Os ON. Os! Oye (OKO 
IIl/19-25 Omer On sO Om Or Or OS Ole" Oa 0 
Ill /26-IV/1 ee Ome iW. One HOeE. (KOLPSO4 wm EOKe. WII Outs 10 
IV/2-8 Me flere Ov ol 24 Oy 0S) Sle, ) Aes pO wean ue ag 
IV/9-15 HOR cOne Ole = OP On CI Ole ew Woes Oye 20 
IV /16-22 CMR OF (Ole Or On 0a ee 0 | ON On 
IV /23-29 CRORE OOM TOM LOR Hier 200.41) IOS a0) ct HO 
IV/30-V/6 CRORE Ones Oiwea Or 1k ON Oke Or 5, 08. 1Ort4 Onl 
Ny 7-13 ONO TOL Orr. 0), 405, 0h | Or le @O., Ve ee 
V/14-20 CMR OR 00 Ont OF 2 ee ok or 2c Beg Sy Ea 
V /21-27 SOMMIO TREE AN) AISait) OLT 38ers) Wee! Bela teO 
V/28-VI1/3 ETM oe Bie Oe 6 Over 1Oin ft Guncstlem, 1:Gen Vila al Qcran 6 
VI/4-10 PMG Ag Oe A olen Baw Dee Bue NO yeni pany 2 
VI/11-17 Ome ae OM) One Sp! Sale g oh tos) QIN scat ae 
VI/18-24 EME SR OL lee: Oar Ane On clay ee’ “Oleg Oe yO 
a een Me ATG on Oe Os 9/4.) On 2 Os Oe a ey iy gh 
VII/2-8 CE Os 2 On On 0s. Or ak Bal nO een 60. 
VII/9-15 oe Oy Or os eh, Oe 5 ee I CO aco 
VII/16-22 CR ee, Th MROE LiL Len RON. Ovi t Caw TO: CAO: TOKO). EO 
VII/23-29 CE ee I Ae Ob Aue IA AOL Oy OF ui cORm. sun nS 
eco Mi cemloe 18; 0; OF ..0 bb *'O. & L 0 OO v4 
VIII/6-12 OO ly Oy On SO I On Od 0, G0 
VIIL/13-19 COTO 1100 SO OVin Omit tii 0) “Oe OOK Ha 
VIII /20-26 OY 020 One: Ole OM 0% ey OF ©. Qoiia-'On, Ma 
Weer 0. . O. 0 0 0, 0 -0 1° 0: O 4 
IX /3-9 CO Or FO | Oto Om 0 OO. OO 
IX /10-16 Ope OR MeIOl TOM N20 WYeOtea TOR. clomihOs MO 4G OCHO 
IX /17-23 OR OM LOL Olas OME Oi tO, ApeO i Out. On Onna 
IX /24-30 See MOM sw um Shs eOrt tt Ot SQ Oi 7! OOOO 
ez MEMO POO) LOTS VOr SOLO OO 0.) O04) O 
X/8-14 DORs Oe Oy MiNi MeO MELONLN0. .0,, Beh. -0.. 1 
X/15-21 (Oe O cunt Oke, Ope in 0; ir 8h isles sO oe 0 
X /22-28 OMe) FLY age ON Os EO ke 2 8 rey AO 
X/29-X1/4 CMe ROMAN ORY 10) MeOMige le) OL TON xO 4S OC WO! APO 
a= 11 CREO, ee Tm OMINOe ) Op 4.0i), 1-0... 0 2 10.0 
XII/17 ye a Me ny ee 
Total A OOO Ae WAY 18) UShue 56) 26 428,,. 48438 - 57-624 


Because of missed days and less than two hour watch, data for 1972 
are not included. That is most unfortunate, because that season fol- 
lowed the coldest winter during the count years. 

The year 1978 was unusual. Following the drought of 1976 and 1977, 


278 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


TABLE 3. Papilio rutulus two hour daily count per week for twelve years at an 
observation point in Citrus Heights, California, U.S.A. 


Date 1970 1971 1973 1974 1975 1976 1977 1978 1979 1980 1981 1982 
III/12-18 i (MM ero 1 © © 2 
III/19-25 i co Pewee OO g 2 
Il/26-IV/1 8. 920° or 90> 26° 71 le 2) oe 
IV/2-8 16°00 P.O FO 1 8 $1 aT) 
IV/9-15 Woe 8 Fe Sb Oe Pe 11m 6? oa 
IV /16-22 i (i i iii mene 6 § 12 Le 
IV /23-29 162 10 0 «3..° 9’ 995.10) WO Beeman 
IV /30-V/6 oY 9.0) 86 11°>. 8 48) (7 6) 4 eee) 
Wels Ci Ty eee ene 2 1G 
V /14-20 Il U5. Be BO 8. 20! a 7 CS ee 
V /21-27 33° 7 2 915’ 65 12 Oo! «6 4 IO oom 
V /28-V1/3 30 5 0: 0 “4.10 74 Shes sts 
VI/4-10 4 14 8 22) 8 5) at) Te! 
VI/11-17 4 6 2) 2) 98 1 8) SOR inane 
VI/18-24 4° 2° OO 20!) (0° 4.0) oF ci one 
VI/25-VIlI/1 15 3 Bh TO 418) a a ee 
VII/2-8 ee. © Ss 8 
VII/9-15 40 2 14 (12 5 199)! 10) “47 (99) os someon 
VII/16-22 97:98 138 9° 16 83° 16 51> 3eSae iia mene 
VII/23-29 50 25. 30 38 ‘30 53. 27 76. AS is arcspemon 
VII/30=VIII/5 28 44 39 44° 40 80 12 58° “Giletgo auoGummo 
VIII /6-12 94°98 139 58 31 17 26) (86> SO4MeiigoramCmmmac 
VIII/13-19 7 20 “Rh AP 38 23 1) 14 Seo RG wanes 
VIII /20-26 5 4 9) 3° 19 JO. 46) 7 Sl fone osama 
VIlL/27-1X/2 15 8 OC DB eG Si ees 
IX /3-9 10 <0 2 “Ys. (5 oe S08) re 
IX/10-16 tt eek 2 OQ FB 
IX /17-23 tO 0 0 0 °O 20 oi BOP Geen 
1X /24-30 Tt i ni) Oo © 8 
X/24 0 0 6-0 0 -0 “0.20 FO ORaOmmENe 
Total 435 206 169 232 234 323 210 198 297 203 306 572 


D. plexippus reached the highest count observed, and P. rutulus reached 
the fourth highest weekly count. Several other species not systemati- 
cally counted were also present in the greatest numbers seen. 

A scrutiny of Tables 1 through 8 will give an understanding of the 
variation of population of the species. It has been said that butterfly 
populations are declining (Moucha, 1974; Newsom-Brighton, 1982). 
These data do not support that statement, but perhaps twelve years’ 
count is too short to say. 

Citrus Heights is not a very good site for counting D. plexippus 
(Table 1), because there are no known Asclepias within miles. How- 
ever, it is in the migratory path to the coastal hibernal colonies. The 
majority of those caught were females, many of which were fluttering 
over vegetation before late August. After that practically all flew straight 
through in a south or southwesterly direction. 


VOLUME 87, NUMBER 4 279 


The data do not show the great swings in D. plexippus population 
reported in the eastern U.S. and Canada (Urquhart, 1960), where in 
some years there are practically none. Locally they varied within more 
narrow limits, from a high count of 147 in 1978 to a low of 83 in 1978. 
There were few spring migrants seen here. The maximum number per 
week was seven in 1977 in the spring. 

D. plexippus reaches a peak population in mid-August of most years. 
Do they start migrating at that time? It is known that they start arriving 
at the Richmond colony, the closest one at a distance at 130 km, in 
mid-September (pers. obs.). 

The count of N. antiopa is the most fascinating of the three species. 
It has the greatest variation from a high of 150 in the week of 11 to 
17 June 1970 to zero in 1974 and 1980. The total yearly count goes 
from a maximum of 400 in 1970 to a minimum of eight in 1974. 

This count supports Shapiro’s (Shapiro, 1974) theory that it goes into 
hibernation after the generation of late May and early June, because 
few sightings are made after that. This behavior is a great mystery 
because there is seemingly ample time and food for the production of 
another generation here in late August and September. It would seem 
to be a great opportunity to increase its numbers and thereby, increase 
survival potential. 

Individual N. antiopa can be seen here in late fall and winter months 
on warm sunny days. The earliest was 19 January 1980 and the latest 
was 17 December 1976. It should be noted there were only four sight- 
ings from the week of 27 August to 2 September to 8 to 14 October. 
There were few sightings after that. 

There is a problem with the count in May and June. There is a grove 
of Salix babylonica 80 m east of the observation point where most of 
the sightings were made. There were so many that the same individual 
was likely counted several times. I arbitrarily decided not to count a 
second appearance for two minutes after the first to reduce duplicate 
counts. If an individual was in view for several minutes nothing was 
counted for the two minutes after it disappeared. 

Fortunately, I started counting N. antiopa in 1970 because of the 
unusually large population that year. From casual observation in 1968 
and 1969 there were few in those years. 

P. rutulus was the most numerous during the count period. How- 
ever, the numbers cannot be compared interspecifically because of 
behavioral differences among the species (specifically, male patrolling 
behavior or territoriality in P. rutulus which would tend to increase 
repeat sightings and hence the count). Once again, only an intraspecific 
index of abundance can be inferred. P. rutulus reached a peak annual 
count in 1982 of 572, followed by another peak of 435 in 1970. The 
minimum annual count was 169 in 1973. It is the last to appear of the 


280 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


three species and the first to disappear. It never appeared before 20 
March, and several years it appeared on that date. In 1975 it didn’t 
appear till 20 April. The latest it appeared was 22 October 1982. 

In 1978 following the driest winters on record of 1976 and 1977, P. 
rutulus reached the fourth highest weekly count of 76 the week of 
23-29 July. In 1982 following the wettest winter this century the three 
consecutive weekly counts starting 23 July reached the highest of 90, 
92, and 96. These facts lead to the obvious but incongruous hypothesis 
that P. rutulus thrives on extremes of rainfall. A possible explanation 
is that these extremes reduce predators. 

Only in 1970, 1973, and 1982 did the spring count accurately predict 
the summer count. The springs of 1970 and 1982 were the highest of 
the twelve (12) years and were followed by high summer counts. The 
spring 1973 count was the lowest followed by the lowest summer count. 
The spring 1978 count was the second lowest followed by the fourth 
highest summer count. Therefore, generally, spring counts are unreli- 
able predictors of summer counts. 

Finally, from these data the July 4th butterfly census of the Xerces 
Society is conducted when few individuals of the species studied are 
on the wing in lowland central California at least, and therefore, po- 
tentially misleading. 


ACKNOWLEDGMENT 


I thank A. M. Shapiro of University of California, Davis for reviewing the manuscript 
and making helpful suggestions for changes. 


LITERATURE CITED 


FIGGINs, W. E. 1971. Climate of Sacramento, California. NOAA Technical Memoran- 
dum NWS-WER-65. Salt Lake City, Utah. 63 pp. 

Moucua, J. 1974. A Color Guide to Familiar Butterflies. Octopus Books Ltd., London. 
190 pp. 

NEWSOM-BRIGHTON, M. 1982. Butterflies are free. National Wildlife 20:27-34. 

SHAPIRO, A. M. 1974. The butterfly fauna of the Sacramento Valley, California. J. Res. 
Lepid. 13:738-82, 115-122, 137-148. 

1979. Weather and the lability of breeding populations of the checkered white 
butterfly, Pieris protodice Boisduval and LeConte. J. Res. Lepid. 17:1-23. 

URQUHART, F. A. 1960. The Monarch Butterfly. University of Toronto Press. Toronto, 
Canada. 361 pp. 


Journal of the Lepidopterists’ Society 
$7(4), 1983, 281-288 


SEASONAL PHENOLOGY OF BATTUS PHILENOR (L.) 
(PAPILIONIDAE) IN CALIFORNIA! 


S. R. Sims? AND A. M. SHAPIRO 


Department of Entomology, University of California, 
Davis, California 95616 


ABSTRACT. The pipevine swallowtail butterfly, Battus philenor (L.), has a flight 
season extending over more than nine months (February to November) in central Cali- 
fornia. The major flight occurs primarily in April and is derived from overwintering 
pupae in diapause. This flight is followed by a partial second generation, consisting of 
9-39% non-diapause first generation offspring. A subsequent temporally scattered flight, 
representing a third generation, is partially derived from earlier season diapausers emerg- 
ing in summer and fall. Under field conditions, pupal diapause intensity progressively 
declines through fall and early winter. Pupal photoperiod response and diapause end by 
mid-winter. There is no sex ratio distortion in either second brood or summer-fall emer- 
gers. Spring field emergence of males tends to precede females, suggesting differences 
in relative rates of post-diapause development. 


A necessary step towards an understanding of the population dy- 
namics and distribution of Lepidoptera is to examine their seasonal 
phenology or timing of recurring periods of activity and dormancy in 
relation to key environmental factors. For species with a diapause phase, 
the appropriate timing of the onset, maintenance, and termination of 
diapause, followed by postdiapause development and resumption of 
reproductive activity is vital to the successful adaptation to their en- 
vironment. 

The pipevine swallowtail, Battus philenor (L.), has an extended flight 
season in central California with adult activity recorded from February 
to November (Opler & Langston, 1968; Shapiro, 1974). Little, however, 
is known about how the flight season is related to the population dy- 
namics and pupal diapause of this species. Shapiro (1975) suggested 
that the long flight season and apparent multivoltinism results from 
reproduction by a non-diapause fraction of each generation. Thus, each 
generation may be a mixture of both continuous developers and indi- 
viduals that undergo an aestivo-hibernal pupal diapause (Masaki, 1980) 
and emerge the following spring. 

In this paper we examine the phenological “strategy” or timing of 
the active and diapause states of philenor as they relate to the annual 
periodicity and variability of the central California habitats. Specifi- 
cally, we estimate the number, timing, and derivation of annual broods 
and determine when diapause terminates under field conditions. 


1 Florida Agricultural Experiment Station Journal Series No. 5026. 
2 Present address: University of Florida AREC, 18905 SW 280th St., Homestead, Florida 33031. 


282 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


SACRAMENTO 1974 


@ MALES 
CO FEMALES 


NUMBER ADULTS EMERGING 


VACA MTNS. 1977 


MARCH APRIL 


Fic. 1. Spring emergence of first brood adult B. philenor from overwintering pupae. 


MATERIALS AND METHODS 


We collected living material from the following populations: Chico, 
Butte Co., CA, 39°42’N (latitude), 60 m (altitude); Davis, Yolo Co., CA, 
38°30'N, 16 m; Vaca Mountains (from Mix Canyon to Solano Lake, 
approx. 7 km SW Winters), Inner Coast Range, Solano Co., CA, 38°25’N, 
50-300 m; Sacramento, Sacramento Co., CA, 38°30/N, 15 m. 

To estimate diapause incidence and adult emergence under field 
conditions, we sampled first generation final instar larvae and prepupae 
from the Davis, Chico, Vaca Mtns., and Sacramento populations during 
mid-May to mid-June from 1974-1976. Samples were maintained out- 
doors, at Davis, on cuttings of the foodplant, Aristolochia californica 
Torr., in large (46 cm side) screened cages. Completion of feeding and 
pupation occurred within one week of collection. Both larvae and pu- 
pae were sheltered from rain but exposed to normal seasonal variations 
of temperature and photoperiod. Individuals were labelled with col- 


VOLUME 387, NUMBER 4 283 


TABLE 1. Percent non-diapause of first brood Battus philenor (A) and percent of 
adults emerging in July, August, or September from first brood diapause pupae >830 days 
old (B). Number in parentheses is sample size; NS = not sampled. 


1974 1975 1976 
A B A B A B 
Davis 25.0 (4) — 23.1 (26) — NS — 
Vaca Mtns. 39.0(141) 3.2 (93) 27.4 (168) — 15:8 (146) 9.7 (123) 
Chico NS — 8.8 (616) 3.7 (135) 14.2 (558) 21.3 (230) 


lection and pupation date and monitored daily for adult emergence 
and sex. We used the same collection, storage, and recording methods 
to study the timing of spring adult emergence the following year. 
The termination of diapause, or completion of diapause “develop- 
ment” (Beck, 1980), can be studied in photosensitive species by deter- 
mining the date at which photoperiod no longer influences the rate of 
morphogenesis; i.e., when morphogenesis is primarily a function of 
temperature and proceeds at a rate similar to that of non-diapause 
individuals (Tauber & Tauber, 1976). Since philenor pupae are pho- 
tosensitive, with the rate of diapause development inhibited under 
short-day photoperiods and increased under long-day photoperiods 
(Sims & Shapiro, 1983), we were able to study field diapause devel- 
opment as follows: Pupae, derived from final instar Chico larvae col- 
lected on 11 June 1976, were maintained outdoors through summer 
and fall in Davis. At monthly intervals from 20 November 1976 to 20 
February 1977, samples of pupae were transferred from their outdoor 
location to LD10:14, LD15:9, and a natural photoperiod (greenhouse) 
at approx. 23.5°C and monitored for emergence. Data were analyzed 
using ANOVA procedures and Duncan’s Multiple Range Test (DMR) 
for significance of differences between means (Sokal & Rohlf, 1969). 


RESULTS 


Our 1974-1977 adult emergence data from overwintered pupae show 
a unimodal peak of spring emergence during the second and third 
weeks of April (Fig. 1). This peak coincides with a period of rapid 
Aristolochia growth. 

Non-diapause pupae and resulting adults obtained from field-col- 
lected first generation larvae provided an estimate of the magnitude 
of the second generation. The second generation ranged from less than 
10% to almost 40% of the surviving first generation pupae (Table 1). 
The Vaca Mtns. population (1974-1976) illustrates that considerable 
yearly variation may occur in second generation size. Mortality of first 
generation pupae was low, ranging from 5-10%. 


284 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


PERCENT EMERGENCE 


3] 3 6 9 12 I5 
MAY JUNE 
PUPATION DATE 


Fic. 2. Mean pupation date and percent adult emergence of B. philenor. Vaca Mtns. 
population; spring 1976. Numbers indicate sample size. 


Pupae from the Vaca Mtns., over 18 days in late spring, 1976, showed 
a marked decrease in adult emergence with increasing pupation date 
(Fig. 2). Increasing diapause was not correlated with temperature (dai- 
ly mean or minimum) but may have been related to host-plant quality 
which deteriorated visibly (increased leaf toughness and decreased suc- 
culence) during this period. A seasonal increase in pupal diapause is a 
possible bias in our estimation of second brood magnitude despite using 
larvae collected from mid-May to early June. Thus, samples taken 
relatively early in the season might overestimate, while late-season 
samples may underestimate non-diapause among first-brood individ- 
uals. 

From first generation pupae in aestival diapause (>80 days post- 
pupation), a small number of adults continued to emerge in July, Au- 
gust, and September (Table 1). The extent to which this scattered flight 
is augmented by a third generation is unknown, but since relatively 
few larvae were observed on Aristolochia after June, the third and any 
subsequent broods are presumably quite small. 

Diapause in the field was considered terminated when there was no 
significant difference in time (days) to adult emergence among the 
short-day, long-day, and natural photoperiods at 23.5°C to which pupae 
were transferred at monthly intervals from November to February 
(Tauber & Tauber, 1976). Using this criterion, pupal diapause ends 


VOLUME 37, NUMBER 4 285 


120 


100 


Fs LD 10:14 
Ua LD 15:9 


DAYS TO EMERGENCE MEAN #ISE 
(ep) 
oO 


20JAN 20FEB 


B0NOV 2ODEC 
SAMPLE DATE 1976-77 


Fic. 8. Adult emergence from diapause B. philenor pupae maintained outdoors prior 
to 23.5°C exposure at the indicated photoperiod. 


during the 30-day period following the winter solstice (Fig. 3). Under 
each photoperiod, adult emergence time was significantly less in Jan- 
uary than in December samples, while emergence times for January 
and February were similar (DMR test, P < 0.05). Pupae from the No- 
vember and December samples displayed a distinct photoresponse. 
November pupae at LD15:9 emerged significantly sooner than those 
at LD10:14 or natural photoperiod, while December pupae at LD15: 
9 emerged sooner than those under a natural photoperiod. 

There was little departure from a 1:1 sex ratio among either second 
brood (51.5% 66, n = 264) or summer-fall emerging individuals (51.6% 
66, n = 64). A comparison of cumulative numbers of males and females 
present after the onset of seasonal emergence shows that males tend to 
emerge somewhat before females (Fig. 4). 


DISCUSSION 


Central California populations of philenor have two major flights 
each year. The first and largest flight occurs in March and April and 
derives from the unimodal emergence of adults from overwintered 
pupae. Most first brood pupae undergo an aestivo-hibernal diapause 
(Masaki, 1980) and overwinter. Some individuals emerge to form a 
partial second generation, the magnitude of which shows both inter- 
and intrapopulation variation (Table 1). There is evidence suggesting 


286 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


200 


FEMALES 


ws 
oe 


175 


I50 


25 


Keye) 


NUMBER OF ADULTS (CUMULATIVE) 


O 2) Ke) I5 20 25 
DAYS AFTER FIRST EMERGENCE 


Fic. 4. Cumulative emergence of spring generation B. philenor adults from over- 
wintering pupae, by sex. Chico, CA population; combined emergence of 1975 and 1977. 


a positive relationship between the pupation date of first brood larvae 
in the field and diapausing frequency (Fig. 2). The increase in diapause 
parallels the visually-determined seasonal decrease in young succulent 
Aristolochia foliage available for larval consumption. Rausher (1981) 
documented a similar seasonal decrease in the quality of Aristolochia 
from Texas. In Texas, the increase in the sclerophyllization (=increased 
leaf toughness and decreased nitrogen) of A. reticulata (Nutt.) between 
March and May led to poorer larval growth and increased larval dis- 
persal of philenor. 

Emergence of adults from the diapausing first generation age-cohort 
is divided between summer-fall and the following spring. The summer- 
fall emergers contribute to the scattered flight of philenor from July 
to November in California. The emergence polymodality represented 
by the aestivo and aestivo-hibernal diapausers of the first generation 
philenor age-cohort does not conveniently fit into any of the phenolog- 
ical categories suggested by Waldbauer (1978). Despite this, it is not a 


VOLUME 37, NUMBER 4 287 


unique example. Among other papilionids, Papilio maacki Fenton 
(Ichinosé, 1974), Eurytides marcellus (Cramer) (Scudder, 1889), and 
some populations of Papilio machaon L. (Wiltshire, 1957) show similar 
discontinuity in pupal diapause duration as does the noctuid moth, 
Barathra brassicae L. (Masaki, 1956; Dolidze cited in Danilevskii, 1965). 
The reproductive success of philenor adults emerging in summer-fall 
is unknown. Since females oviposit exclusively on tender growing shoots 
and first instar larvae can only feed on these, opportunities for summer 
reproduction are usually limited. Following the onset of fall rains in 
September and October, a small amount of new Aristolochia growth 
may become available, and we have found both ova and final instar 
larvae on this growth in late October. 

Diapause development in philenor is completed before midwinter 
(Fig. 3). No photoresponse was found in pupae sampled one month 
after the winter solstice; the development rate of pupae at this time 
was similar to non-diapause pupae at the same temperature. It is most 
likely that morphogenesis and adult emergence in the spring following 
diapause termination are functions of temperature accumulations above 
a minimum temperature developmental threshold. 

Previous observations that individuals of later broods are primarily 
males (Fee, 1979) are not supported by our results which show no sex- 
ratio distortion among either second brood or summer-fall emergers. 
The only evidence for sex-related phenological differences was ob- 
tained from adult emergence from overwintering post-diapause pupae. 
Females lag slightly behind males in emergence times, suggesting a 
greater heat unit requirement among females for completion of de- 
velopment. 


ACKNOWLEDGMENTS 


We thank S. O. Mattoon for field assistance, J. L. Hatfield for climatic data, C. F. 
Satterwhite for skillful manuscript preparation, B. Blau and M. Tatar for discussion and 
advice. 


LITERATURE CITED 


BECK, S. D. 1980. Insect Photoperiodism. Second ed. Academic Press, NY. 387 pp. 

DANILEVSKII, A. S. 1965. Photoperiodism and Seasonal Development of Insects. Oliver 
& Boyd, Ltd., London. 283 pp. 

FEE, F. D. 1979. Notes on the biology of Battus philenor (Papilionidae) in Centre 
County, Pennsylvania. J. Lepid. Soc. 33:267-268. 

ICHINOSE, T. 1974. Pupal diapause in some Japanese papilionid butterflies, with special 
reference to the difference in photoperiodic response between the diapausing pupae 
of Papilio maackii Fenton and P. xuthus Linnaeus. Kontyu 42:439-—450. 

MasakI, S. 1956. The local variation in the diapause pattern of the cabbage moth, 
Barathra brassicae Linné, with particular reference to the aestival diapause (Lepi- 
doptera: Noctuidae). Bull. Fac. Agric. Mie Univ. 13:29-46. 

1980. Summer diapause. Annu. Rev. Entomol. 25:1-25. 


288 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


OpLER, P. A. & R. L. LANGSTON. 1968. A distributional analysis of the butterflies of 
Contra Costa County, California. J. Lepid. Soc. 22:89-107. 

RAUSHER, M. D. 1981. Host plant selection by Battus philenor butterflies—the roles of 
predation, nutrition, and plant chemistry. Ecol. Monogr. 51:1-20. 

SCUDDER, S. H. 1889. The butterflies of the eastern United States and Canada. Vol. II. 
Cambridge, MA. 

SHAPIRO, A. M. 1974. The butterfly fauna of the Sacramento Valley, California. J. Res. 
Lepid. 13:73-82, 115-122, 1387-148. 

1975. The temporal component of butterfly species diversity. Pp. 181-195, in 
M. L. Cody & J. M. Diamond (eds.). Ecology and Evolution of Communities. Belknap 
Press, Cambridge, MA. 

Sims, S. R. & A. M. SHAPIRO. 1983. Pupal diapause in Battus philenor (L.) (Lepidoptera: 
Papilionidae). Ann. Entomol. Soc. Amer. 76:407-412. 

SOKAL, R. R. & F. J. ROHLF. 1969. Biometry. W. H. Freeman, San Francisco. 776 pp. 

TAUBER, M. J. & C. A. TAUBER. 1976. Insect seasonality: Diapause maintenance, ter- 
mination, and postdiapause development. Annu. Rev. Entomol. 21:81—107. 

WALDBAUER, G. P. 1978. Phenological adaptation and the polymodal emergence pat- 
terns of insects. Pp. 127-144, in H. Dingle (ed.). The Evolution of Insect Migration 
and Diapause. Springer-Verlag, NY. 

WILTSHIRE, E. P. 1957. The natural history of Papilio machaon L. in Bagdad. Trans. 
R. Entomol. Soc. Lond. 110:221-248. 


Journal of the Lepidopterists’ Society 
37(4), 1983, 289-300 


DISTRIBUTION AND NOTES ON THE GREAT DISMAL 
SWAMP POPULATION OF MITOURA HESSELI RAWSON 
AND ZIEGLER (LYCAENIDAE) 


ANDREW F. BECK 


Department of Entomology, Virginia Polytechnic Institute and State University, 
Blacksburg, Virginia 24061 


AND 


WILLIAM J. GARNETT 
314 Reynolds Street, Apt. C, Blacksburg, Virginia 24060 


ABSTRACT. The microdistribution of M. hesseli within selected areas of the Dismal 
Swamp (VA and NC) is found to be coincident with the occurrence of its larval foodplant, 
Chamaecyparis thyoides (L.) B.S.P. (Cupressaceae). Observations on nectar feeding, ap- 
parent predation on adults by birds, perching behavior by adult males, and other behav- 
ioral phenomena are reported. Two new categories of beak-inflicted wing damage in 
Lepidoptera are described, and a possible selective advantage for the dorsal “false head” 
found in many lycaenid species is discussed. The white spot of the discal cell of the 
ventral forewing is found to be an unreliable character for separating M. hesseli from 
M. gryneus (Hiibner) in Virginia, but the subterminal brown bars in cells M, and M, of 
the ventral hindwing are unique to M. hesseli. 


Since its original description and the subsequent description of its 
early stages (Rawson et al., 1951), little has been published concerning 
the biology or behavior of Hessel’s hairstreak, Mitoura hesseli Rawson 
and Ziegler (1950). Progressive range extensions have been reported 
(Pease, 1963; Anderson, 1974; Johnson, 1978; Baggett, 1982); and it 
appears that this insect will be found throughout the range of its larval 
foodplant, Chamaecyparis thyoides (L.) B.S.P. (Cupressaceae). 

The geographic proximity and the morphological and biological sim- 
ilarities between M. hesseli and M. gryneus (Hiibner) suggest recent 
speciation. Although the normal foodplant for M. gryneus is Juniperus 
virginiana L. (Cupressaceae), it has been successfully reared on C. 
thyoides (Remington & Pease, 1955); and Gifford and Opler (1983) 
have reared M. hesseli on J. virginiana. The wing patterns of the two 
species are nearly identical, and the genitalic similarities (and differ- 
ences) were reported by Johnson (1976). 

In view of this close biological relationship, the reported behavioral 
differences between the two species appear striking. The literature 
suggests that, except for at the type locality, M. hesseli is an infrequent 
find even in the vicinity of C. thyoides and is best collected at flowers 
near the foodplant rather than on the foodplant itself. The pugnacious 
territoriality of adult male M. gryneus is well known (Johnson & Borgo, 
1976), and experience with this species in Virginia shows that it is 


290 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


rather ubiquitous. A short hedgerow of several J. virginiana is ade- 
quate to support a double-brooded colony. It is intriguing that the 
microdistribution and adult behavior of M. hesseli should vary so greatly 
from its closest extant relative. The authors were able to study the M. 
hesseli population in the Great Dismal Swamp National Wildlife Ref- 
uge, located in southeastern Virginia and northeastern North Carolina, 
for the purpose of clarifying the nature of these differences and per- 
haps uncovering some explanation for them. 


MATERIALS AND METHODS 


The study site was the Great Dismal Swamp National Wildlife Ref- 
uge (the Refuge), located approximately between latitudes 36°26’N and 
36°48’N and longitudes 76°22’W and 76°33’W. A thorough character- 
ization of the Refuge and surrounding swamp was given by Kirk (1979). 
Chamaecyparis thyoides is found in the Refuge as an invader of ditch 
edges, as a member of variable dominance in a generally mixed hard- 
wood forest, and in pure stands of many hectares extent (Fig. 1). Roads 
and ditches provide the only access to the Refuge interior, although it 
is possible to penetrate off-road areas on foot with great difficulty. All 
roads follow ditches, but many ditches are unaccompanied by roads 
and are often impassable due to rooted and fallen vegetation. 

Several trips were made in 1981 to scout potential sites for locating 
M. hesseli. With the aid of a vegetation map provided by the Refuge 
administration, those areas of C. thyoides accessible by vehicle were 
identified. In 1982, a qualitative sampling program was begun. Select- 
ed 0.8 km (0.5 mi.) sections along passable roads were sampled for M. 
hesseli. Each section was sampled at least once, and there was no 
uniformity of sampling effort. With the one exception described below, 
the collection or positive sight identification of two specimens was 
sufficient to consider a section positive for M. hesseli. Flowering shrubs, 
vegetation perches, and damp patches in the road were examined thor- 
oughly. Enough other spring species were in flight to ensure that sec- 
tions not near C. thyoides would be examined as closely as those near 
the foodplant. This regimen was followed on 3, 18, 19, and 20 April 
and in the late afternoon only of 2 April. Approximately 24 km of road 
were examined in this manner, and sections were selected so that about 
one third were in areas where C. thyoides could be seen along the 
road or in the forest. The remaining sections were at various distances 
from the foodplant. 

On 18 April, a 5- to 6-meter-wide trail through a dense stand of 
mature C. thyoides was discovered and followed for ca. 1 km (Fig. 1, 
point A). The edges of the trail were lined with immature C. thyoides, 


VOLUME 37, NUMBER 4 291 


LAKE 
| DRUMMON 


4miles 


3 


fo} 1 2 3 4 5 kitometers 


RANGE OF C. THYOIDES 


DITCHES WITH ROADS Saas 


DITCHES WITHOUT ROADS --=—- 


Fic. 1. Distribution of Chamaecyparis thyoides within the Great Dismal Swamp 
National Wildlife Refuge in relation to roads and ditches. Point A is site of trail discussed 
in text. (Adapted from U.S. Geological Survey open-file map 76-615.) 


292 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


4-6 m tall, behind which loomed the crowns of mature trees, 18-22 
m tall. Mitoura hesseli was abundant along this trail, and a series was 
collected for later examination. The remaining field data consisted of 
general observations on the behavior and habits of this butterfly species. 

Out of the field, 32 adults were sexed and examined for the presence 
of both the white spot in the discal cell of the ventral forewing and 
the brown bars distad of the postmedian line in cells M, and M, of the 
ventral hindwing, several characters used by authors to differentiate 
M. hesseli from M. gryneus (Rawson & Ziegler, 1950; Clench, 1961; 
Howe, 1975). A series of M. gryneus (n = 74) was similarly examined. 
The latter specimens were collected in Virginia, although none was 
collected in the vicinity of the Refuge. Rudimentary white spots of 
only several scales were considered as absent. 

A single visit to the Refuge was made on 6 July 1982 expressly to 
photograph M. hesseli in its natural setting. Some supporting obser- 
vations were made at this time. 


RESULTS AND DISCUSSION 


Distribution. Positive and negative collection sites are indicated in 
Fig. 2. Ditches and roads have been removed from this figure for 
clarity. The negative results in the vicinity of a mature stand of C. 
thyoides (Fig. 2, point A) are likely artifactual. This site was visited at 
ca. 0900 EST on an overcast day with winds to 77 kph (48 mph) and 
was the only site in the vicinity of the foodplant which did not yield 
M. hesseli. Summer-brood individuals were abundant here on 6 July, 
and it is assumed that the aforementioned weather conditions were 
responsible for the negative findings in April. Neighboring areas proved 
to be densely colonized when examined under more favorable weather 
conditions. Point B in Fig. 2, in combination with the positive samples 
to the east of it, suggests that M. hesseli is likely found throughout that 
northwestern stand of C. thyoides. Other such opportunities (in which 
a stand could be bracketed by samples) were unfortunately unavail- 
able. Point C in Fig. 2 is the only section which was positive for M. 
hesseli based only on sight records. No C. thyoides grew along the 
road or in the forest along this section, but it had recently invaded the 
far bank of the adjacent ditch. Mitoura hesseli was seen nectaring on 
blossoms of Vaccinium corymbosum L. (Ericaceae) along that bank, 
just out of reach of our nets. Sights not in the vicinity of C. thyoides 
were consistently negative. 

General observations. Vaccinium corymbosum was the dominant 
flowering plant in the Refuge on 2, 3, and 13 April; and it rarely grew 
far from C. thyoides. In contrast to the findings of Rawson and Ziegler 
(1950) in New Jersey, M. hesseli was found to utilize V. corymbosum 


VOLUME 37, NUMBER 4 293 


RANGE OF GC. THYOIDES 


M. HESSEL! PRESENT @ 


: ABSENT (C) 


Fic. 2. Results of qualitative sampling program, illustrating the distribution of Mi- 
toura hesseli in relation to its larval foodplant, Chamaecyparis thyoides, within the Great 
Dismal Swamp National Wildlife Refuge. Points A and B are discussed in text; Point C 
is based on sight records only. 


294 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


readily as a nectar source. Most specimens thus collected were female, 
but the actual sex ratio was unrecorded. By 19 April, V. corymbosum 
was past flowering; and other plants with varied distributions were 
beginning to flower. Amelanchier intermedia Spach (Rosaceae) was 
the only additional bloom on which M. hesseli was seen to nectar in 
the spring. Sassafras albidum (Nutt.) Nees (Lauraceae) and an un- 
identified willow (Salix sp., Salicaceae) were flowering locally but not 
near any site at which M. hesseli was recorded. Summer-brood indi- 
viduals were seen to utilize Cephalanthus occidentalis L. (Rubiaceae), 
Phytolacca americana L. (Phytolaccaceae), and Apocynum sp. undet. 
(Apocynaceae) as nectar sources. Despite many fresh flower heads of 
Achillea millefolium L. (Compositae) and Daucus carota L. (Umbel- 
liferae) in the immediate vicinity of abundant M. hesseli, neither was 
ever visited by the butterfly during several hours of observation (late 
afternoon, 6 July). 

The flight of M. hesseli when nectaring is very distinct from that of 
M. gryneus. While the latter retains its darting flight when approach- 
ing nectar sources, the former assumes a fluttering, casual flight, at 
least at Vaccinium blossoms. In one instance, while beating bushes to 
dislodge perching or nectaring individuals, a female M. hesseli was 
seen to remain undisturbed even though the flower cluster on which 
she was nectaring was roughly shaken. In July, a second female was 
perched out of camera range on a C. occidentalis blossom, and she 
could not be dislodged with repeated, direct taps of the net handle. 
The senior author broke off the branch on which she was perched and 
brought it into a clear area where he was able to photograph the 
specimen at close range for several minutes until the butterfly, appar- 
ently sated, flew away. 

Two or three specimens of M. hesseli (sex unrecorded) were seen to 
flounder across the road as if in physical distress. When collected, these 
proved to be fresh, post-teneral specimens with no evident, external 
injuries. These may have been diseased or parasitized individuals, or 
they may have been struck by passing vehicles. The latter possibility 
is unlikely since there is almost no vehicular traffic within the Refuge, 
but it cannot be discounted. No attempt was made to culture disease 
agents or rear parasites from these specimens. 

The walk along the trail revealed hitherto unreported behavior pat- 
terns in M. hesseli. Here this species behaved much the same as M. 
gryneus. Males were seen perching on immature C. thyoides and dart- 
ing out after passing butterflies and other insects. Most were seen on 
the sunlit side of the trail, and most selected perching positions in the 
top third of the trees. Numerous “dogfights” were seen involving two 
or three individuals, and individuals were occasionally seen visiting 


VOLUME 37, NUMBER 4 295 


Vaccinium blossoms, at which their flight showed no sign of the le- 
thargic pattern described earlier. No females were seen or collected 
along this trail except for a single specimen collected on blossoms near 
the far end of the trail. No activity was seen around the canopy of the 
surrounding stand of mature foodplant, but the distance precluded 
conclusive observations. Two specimens (sexes unrecorded) were seen 
to land upon the trail and walk about for several cm, eventually climb- 
ing down into crevices formed by fallen limbs in the mud. Here they 
would quietly sit with only the tips of their hindwings exposed. These 
individuals are presumed to have been tippling ground moisture, al- 
though the forward portion of their bodies could not be seen to confirm 
this. It is also not known why they crawled into crevices to get moisture, 
since most of the trail surface was mud. No matings were observed for 
this species. 

Predation. Evidence of predation by birds was seen in specimens 
collected during this study. Although no attacks were observed, nu- 
merous insectivorous birds are found in the area (Anonymous, 1980). 
Since Sargent’s (1976) classification of beak damage was designed for 
and applied to noctuid species, no category (Type I, Type II, or Type 
III) is descriptive of the damage inflicted on Lepidoptera which rest 
with wings folded upright over the back. The resulting damage from 
an attack on an insect in this position is manifested in either two or 
four wings and is always bilaterally symmetrical. In keeping with Sar- 
gent’s (1976) nomenclature, the names Type IV and Type V damage 
are suggested. Type IV damage (Fig. 3) is caused by attacks in which 
the beak is oriented roughly parallel to the major veins in the insect’s 
wings. The beak crosses the wing margin rather than the costa. Three 
subcategories are recognized: IVa, which involves only the forewings; 
IVb, which involves only the hindwings and is typical of thecline ly- 
caenids (Robbins, 1980); and IVc, which involves both fore- and hind- 
wings. Type V damage (Fig. 4) is caused by bites which cross the 
forewing costa. Here two subcategories are possible: Va, which does 
not extend to the hindwing costa; and Vb, which does. 

Type IV damage generally results in notches in the wings, but Type 
V damage rarely does. In the latter case, beak imprints instead of 
notches are left on the forewing (in the case of Type Va attack), and 
no example of Type Vb damage has been seen. It is likely that Type 
Vb attacks are almost always successful due to the unlikelihood of the 
forewing costa tearing to allow the insect to escape (Robbins, 1980). 
This leaves to be explained why individuals with Type Va damage are 
observed, since the predator grasps the prey by the costae in this type 
of attack, also. The authors suggest that the insect’s reaction to a Type 
Va attack is to snap open the hindwings (as a natural attempt at flight), 


296 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Fic. 3. Diagrammatic illustration of Type IV bird attack and resulting wing damage. 
A) Type IVa involves forewings only; B) Type IVb involving hindwings only (illustrated 
with remaining anal fragments removed, as is the case in many field-collected specimens); 
C) Type IVc involving all wings (after Sargent, 1976). 


startling the bird and thus facilitating an escape. This may help to 
explain why so many thecline species exhibit a rudimentary “false 
head” on the dorsum of the hindwings as well as the more well de- 
veloped one on the ventrum. Besides providing a deflection target for 


VOLUME 37, NUMBER 4 297 


Fic. 4. Diagrammatic illustration of Type V bird attack and resulting wing damage. 
A) Type Va involving only the forewings; B) Type Vb involving all wings (after Sargent, 
1976). 


Type I attacks (attacks while in flight), an eyespot in this position would 
contribute to the startle effect in the event of a Type Va attack. 

Fig. 5 illustrates damage types IVb, IVc, and Va, as found in the 
Refuge population of M. hesseli. 

Wing maculation. The 15 male M. hesseli examined showed no 
white spot in the discal cell of the ventral forewing. Fifteen of 17 
females (88.2%) had the white spot; the remaining two did not. Overall, 
46.9% (n = 32) had the white spot. Of 74 M. gryneus examined, 6 of 
58 males (10.3%) and O of 16 females had the spot. Overall, 8.1% had 
the spot. 

This character does not appear to be reliable enough for field iden- 
tification and should probably be omitted from future keys separating 
these two species. This trait appears sex dependent, reversed from one 
species to the other, but the significance of that (or even its validity) 
is uncertain based on these limited data. 

All M. hesseli and no M. gryneus examined showed the brown bars 


298 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


Fic. 5. Mitoura hesseli showing Types IV and V wing damage. A) Type IVb damage 
(6, Great Dismal Swamp, Camden Co., NC, 13-iv-1982):; B) Type IVc damage (6, same 
data as A) with anal area of left hindwing missing, also; C) Type Va damage with beak 
imprints indicated by arrows (2, Great Dismal Swamp, Suffolk, VA, 13-iv-1982). 


VOLUME 37, NUMBER 4 299 


distal to the postmedian line in cells M, and M, of the ventral hind- 
wings. The number examined was slightly less for both species because 
of individuals with Type IVb damage which obliterated this character. 
We suggest that this character be used for field separation of these two 
species. 


SUMMARY 


The Great Dismal Swamp harbors a large population of Mitoura 
hesseli Rawson and Ziegler. With one exception which may be ex- 
plained by poor weather conditions at the time of the spring visit, all 
_ sample areas containing Chamaecyparis thyoides (L.) B.S.P., produced 
Hessel’s hairstreak. Summer-brood individuals were abundant at this 
one negative site. Areas narrowly removed from the foodplant were 
consistently nonproductive. Vaccinium corymbosum L., Cephalanthus 
occidentalis L., Phytolacca americana L., Apocynum sp., and Ame- 
lanchier intermedia Spach were observed as nectar sources for M. 
hesseli, and females were more common at flowers than were males. 
An area of immature C. thyoides at the margin of a mature, pure 
stand of that species revealed M. hesseli males perching and darting 
in a manner indistinguishable from M. gryneus (Hiibner). No matings 
were observed, and it is suggested that the mature foodplant canopy 
be examined for its role in the ecology of M. hesseli. Evidence of 
predation by birds.was seen in many collected specimens, and new 
categories of wing damage, Types IV (with three subcategories) and 
V (with two subcategories), are proposed to accommodate damage in 
species holding their wings folded upright at rest. It is suggested that 
the rudimentary, dorsal “false head” found in certain Lycaenidae may 
provide protection against certain kinds of predator attack. The white 
dot in the discal cell of the ventral forewing was found to be an un- 
reliable field character for separating M. hesseli from M. gryneus. The 
character is possibly sex linked although linked to opposite sexes in 
these two species. It is found in 46.8% of M. hesseli (but never in 
males) and 8.1% of M. gryneus (but never in females). The subterminal 
brown bars in cells M, and M, of the ventral hindwing of M. hesseli 
were found to be reliable characters for separating this species from 
M. gryneus, in which they are absent. 


ACKNOWLEDGMENTS 


Collecting in the Refuge was made possible through the permission of the U.S. De- 
partment of the Interior and the cooperation of the Great Dismal Swamp National 
Wildlife Refuge management, Suffolk, VA. Dr. E. C. Turner, Jr. (Department of Ento- 
mology, VPI & SU, Blacksburg, VA) made transportation available; and the administra- 
tion of the Tidewater Research Station, VPI & SU, Holland, VA, provided a much needed 
place to spend the nights. We are indebted to Dr. T. D. Sargent and Dr. R. K. Robbins 
for their helpful comments on an early draft of this manuscript. 


300 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


LITERATURE CITED 


ANDERSON, RICHARD A. 1974. Southern records of Mitoura hesseli (Lycaenidae). J. 
Lepid. Soc. 28(2):161-162. 

ANONYMOuS. 1980. Birds of Back Bay. U.S. Dept. Inter. FWS. publ. no. RL51510-02. 

BAGGETT, H. DAviID. 1982. Season summary. News Lepid. Soc. No. 6 (Nov./Dec.). 

CLENCH, HARRY K. 1961. Tribe Theclini. Pp. 206-207, in Paul R. Ehrlich & Anne H. 
Ehrlich. How to Know the Butterflies. Wm. C. Brown Co., Dubuque, Iowa. 262 pp. 

GIFFORD, SAMUEL M. & PAUL A. OPLER. 1983. Natural history of seven hairstreaks in 
coastal North Carolina. J. Lepid. Soc. 37(2):97-105. 

Howe, WILLIAM H. 1975. The Butterflies of North America. Doubleday and Co., 
Garden City, NY. 633 pp. + 97 plates. 

JOHNSON, KurRT. 1976. Three new nearctic species of Callophrys (Mitoura), with a 
diagnosis of all nearctic consubgeners (Lepidoptera: Lycaenidae). Bull. Allyn Mus. 
38:1-30. 

1978. Specificity, geographic distributions, and foodplant diversity in four Cal- 
lophrys (Mitoura) (Lycaenidae). J. Lepid. Soc. 32(1):3-19. 

JOHNSON, KURT & PETER M. Borco. 1976. Patterned perching behavior in two Cal- 
lophrys (Mitoura) (Lycaenidae). J. Lepid. Soc. 30(3):169-183. 

KIRK, PAUL W., JR. 1979. The Great Dismal Swamp. Univ. Press of Virginia, Char- 
lottesville, VA. 427 pp. 

PEASE, ROGER W., JR. 1963. Extension of known range of Mitoura hesseli. J. Lepid. 
Soc. 17(1):27. 

RAWSON, GEORGE W. & J. BENJAMIN ZIEGLER. 1950. A new species of Mitoura Scudder 
from the pine barrens of New Jersey. J. N.Y. Entomol. Soc. 58(2):69-82. 

RAWSON, GEORGE W., J. BENJAMIN ZIEGLER & SIDNEY A. HESSEL. 1951. The immature 
stages of Mitoura hesseli Rawson and Ziegler (Lepidoptera, Lycaenidae). Bull. 
Brooklyn Entomol. Soc. 46(5):123-130. 

REMINGTON, CHARLES L. & ROGER W. PEASE, JR. 1955. Studies in foodplant specificity 
1. The suitability of swamp white cedar for Mitoura gryneus (Lycaenidae). Lepid. 
News 9(1):4-6. 

ROBBINS, ROBERT K. 1980. The lycaenid “false head” hypothesis: Historical review 
and quantitative analysis. J. Lepid. Soc. 34(2):194-208. 

SARGENT, THEODORE D. 1976. Legion of Night (the Underwing Moths). Univ. Mass. 
Press, Amherst, MA. 222 pp. 


Journal of the Lepidopterists’ Society 
37(4), 1983, 301-305 


A NEW SPECIES OF SPARTINIPHAGA (NOCTUIDAE) 
FROM THE NEW JERSEY PINE BARRENS 


DALE F. SCHWEITZER 


Peabody Museum, Yale University, P.O. Box 6666, 
170 Whitney Ave., New Haven, Connecticut 06511 


ABSTRACT. Spartiniphaga carterae Schweitzer is described as a new species from 
the New Jersey Pine Barrens. The type series consists of 5 66, 4 92 and 1 other specimen 
is known, all from Burlington County, New Jersey. The species is quite distinctive in 
maculation and male genitalia, but appears to be related to Spartiniphaga inops (Grote). 


The following species is genitalically close to Spartiniphaga inops 
(Grote). So far as known the ten specimens cited below are the only 
ones extant in collections. Nothing is known of its life history. 


Spartiniphaga carterae Schweitzer, new species 


Forewing. Male: Light ochreous, rather uniform. Lines often virtually invisible except 
for series of dark dots on the veins representing the postmedian, and a single such dot 
representing the antemedian. Two paratypes have dark postmedian and subterminal lines 
like S. inops. Reniform with faint dark outer and pale inner rings, broader costad, with 
black spot in anterior end. Orbicular a faint dark circle. Female: All markings tending 
to be lost, the allotype nearly immaculate, but entire pattern seen on males is traceable 
on the paratype. Ventrally powdered with fuscous, no pattern. 

Forewing length: 13.8-15.2 mm 46; 11.7-13.1 mm 99. 


Fic. 1. Spartiniphaga carterae, holotype é. 


302 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Fic. 2. Spartiniphaga carterae, allotype &. 


Hindwing. Very pale with slight ochreous tinge in the males, nearly pure white in the 
females. Unmarked above, except for brown postmedian line on one 6. Ventrally with 
prominent discal spot and fuscous along margins. 

Body. Head and thorax concolorous with forewing, abdomen paler. Thoracic tufts 
weak, abdominal basal tuft vestigial. 

Male genitalia. Illustrated in Figs. 4a and 4b. Differences from S. inops (Figs. 5a, 5b) 
are listed in Table 1. The easiest characters for practical use are the size and shape of 


Fic. 3. Spartiniphaga inops, 6, Mystic, New London Co., Conn., 1 Sept. 1925, leg. 
H. P. Wilhelm (YPM). 


VOLUME 37, NUMBER 4 3038 


Fics. 4a,b & 5a, b. 4a, male genitalia of Spartiniphaga carterae, holotype (aedeagus 
removed)—b, aedeagus, vesica everted; 5a, male genitalia of S. inops from same moth 
as in Fig. 3—b, aedeagus, vesica everted, specimen from Martha’s Vineyard, Mass., 10 
Sept. 1945, leg. F. M. Jones (YPM). 


the basal tooth on the cucullus, which is spine-like on S. carterae, and the size and shape 
of the single large thorn-like spine in the vesica. 

The presence of cucullar teeth on both species, the similarity in valve shape and in 
the vesica suggest that S. carterae and S. inops are fairly close relatives. 


TABLE 1. Comparison of the male genitalia of Spartiniphaga carterae and S. inops. 
Two inops and three carterae were examined in temporary glycerin mounts. 


Character S. carterae S. inops 
Cucullar teeth Basal one long, simple and Four very small, largely 
pointed; 3 or more well fused, basal pair somewhat 
separated smaller teeth set off, larger 
Ampulla Short, slightly notched Longer, simple 
Saccular lobes Somewhat pointed Blunt 
Juxta Deeply cleft Very shallowly cleft! 
Aedeagus 
“Thorn” Long, with well defined Much shorter, evenly tapered 
point to a point 
Cornuti Longer, apparently more nu- Shorter, apparently fewer 
merous 


' The juxta cannot be clearly seen in the figure of S. inops. 


304 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Fics. 6a, b. a, bursa copulatrix of Spartiniphaga carterae, paratype; b, ovipositor and 
associated structures. 


Female genitalia. Illustrated in Figs. 6a and 6b, but not compared with other species. 

Diagnosis, superficial. Spartiniphaga carterae can usually be distinguished from S. 
inops by the greatly reduced forewing pattern. S. inops (Fig. 3) apparently always has 
well defined antemedian, median, postmedian, and subterminal lines. Also the dots rep- 
resenting the postmedian line on the veins are larger on three male S. carterae than on 
most S. inops seen. 

Types. HOLOTYPE: 4, N.J.: Burlington Co., Batsto, 22 September 1973 at UV. leg. 
Dale F. Schweitzer. Illustrated in Fig. 1. Allotype: 9, same data except 21 September 
1975 at MV trap. Paratypes: 6 same data as allotype; 2 same locality, but 19 September 
1970, UV trap, leg. Dale F. Schweitzer and Annie Carter; Whitesbog, N. J., E. P. Dar- 
lington, 17 Sep. 1936 (6, 222), 20 Sep. 1940 (6), 27 Sep. 1938 (68). 

The holotype and allotype are in the type collection of the Peabody Museum of Natural 
History, Yale University (YPM). Batsto paratypes are in my collection. The others are at 
the Carnegie Museum, Pittsburgh, PA and the Montshire Museum, Hanover, N.H. (1938 
3). I have seen one other specimen of this species, collected by John W. Cadbury III and 
in his collection, from Whitesbog, N.J., 12 October 1940. 

Dedication. This species is named for Annie Carter, the naturalist at Batsto Village 
(Wharton State Forest) who tended the trap in which the first Batsto specimen was taken 
and whose hospitality and assistance has contributed immensely to my research on Pine 
Barrens Lepidoptera since 1968. ; 

Distribution. Spartiniphaga carterae is known only from Burlington County, New 
Jersey and is probably restricted to the New Jersey Pine Barrens. 


The Batsto series was collected at the Batsto Nature Center near the 
east bank of the Batsto River, a few hundred meters upstream from 


VOLUME 37, NUMBER 4 305 


the dam at Batsto. The larva probably bores in one of the sedges 
growing in or along the Batsto or Mullica Rivers. No other species of 
Spartiniphaga has been collected in New Jersey. 


ACKNOWLEDGMENTS 


I thank Douglas C. Ferguson for examining and commenting upon one male paratype. 
The authorities at the following institutions permitted me to examine the collections 
under their care: American Museum of Natural History, New York; Carnegie Museum; 
Montshire Museum; Florida Department of Plant Industry; United States National Mu- 
seum. The personal collections of H. D. Baggett, John W. Cadbury III, C. P. Kimball 
and Joseph Muller were also examined. I thank Joseph Gall for the use of his Zeiss 
Photomicroscope with which I took the genitalia photographs. 


Journal of the Lepidopterists’ Society 
37(4), 1983, 306-309 


THE LARVA OF HOMORTHODES FURFURATA 
(GRT.) (NOCTUIDAE) 
KENNETH A. NEIL 


Department of Biological Sciences, Simon Fraser University, 
Burnaby, B.C., Canada V5A 1S6 


ABSTRACT. The mature larva of Homorthodes furfurata (Grt.) is described and 
illustrated. 


Homorthodes furfurata (Hadeninae) was described by A. R. Grote 
in 1874 based on material collected at Albany, New York. To date, 
nothing has been published on the immature stages of this species. 
Crumb (1956) described the larva of what he thought was H. furfurata 
based on an incorrect determination of the adults by Benjamin. As 
shown by Godfrey (1972), Crumb’s determination actually applied to 
the closely related western species H. uniformis (Smith). Rockburne 
and Lafontaine (1976) gave maple (Acer sp.) as the host plant. 

H. furfurata occurs from Nova Scotia (Ferguson, 1954), Maine, Que- 
bec and Ontario, south to Massachusetts, and central New York state 
(Forbes, 1954). A female H. furfurata was collected at an ultraviolet 
light on 29 July 1978, 2.5 km south of Tomahawk Lake, Halifax Coun- 
ty, Nova Scotia, and over the next week laid 10 eggs in a holding 
container. | 

The first instar larvae were confined with both living and dead leaves 
of maple, oak (Quercus sp.), cherry (Prunus sp.), Osmaronia sp., and 
Taraxacum officinale Weber as well as an artificial diet based on that 
of Hinks and Byers (1976). The larvae accepted only living Taraxacum 
officinale leaves but grew slowly with only two reaching maturity. Both 
larvae pupated by 10 October due to the constant conditions of labo- 
ratory rearing. Neither pupa survived the winter. H. furfurata over- 
winters as a larva with adults emerging the following July and August. 

This paper describes the mature larva of H. furfurata. All illustra- 
tions were drawn to scale using a camera lucida and stereomicroscope. 
The terminology and abbreviations used follow Godfrey (1972). 


Homorthodes furfurata (Grote) 


General. Head: integument with minute granules; width 8.0 mm. Total length 25.8 
mm. Body: integument with minute granules; Ab7-8 distinctly swollen; tapering ceph- 
alad. Prolegs present on Ab3-6, size increasing posteriorly; those on Ab3 slightly more 
than ¥% the size of those on Ab6. Crochets uniordinal, 23-25 per third abdominal proleg, 
25-29 per fourth, 28-34 per fifth, 33-37 per sixth. All setae simple. 

Coloration (living material). Head (Fig. 3): blackish brown with a few black coronal 
freckles. Body (Figs. 1, 2): blackish, paler on ventral surface. Middorsal and subdorsal 
lines, whitish, narrow and broken, reduced to a series of dashes. Dorsal and subdorsal 
setal bases whitish. Spiracles dark orange-brown with black peritremes. Lateral shield of 


VOLUME 37, NUMBER 4 307 


a4 Aa A . oh) 4 ik , ry : 
Co ee SFE IPN Teo oy ha? 


ad 


2 


Fics. 1 & 2. Homorthodes furfurata, larva: 1, lateral view; 2, dorsal view (5.5). 


prolegs brownish black. Prothoracic.shield orange-brown with lateral and posterior mar- 
gins black. 

Head (Fig. 3). Epicranial suture 0.45 mm long; height of frons (apex to Fa’s) 0.49 mm; 
distance from F 1 to anterior edge of clypeus 0.37 mm; interspace between F1-F1 0.23- 
0.25 mm; AFa anterior and AF2 posterior to apex of frons; Al-A8 forming an obtuse 
angle at A2; P1-P1 0.62-0.63 mm; P2—P2 0.64-0.65 mm. Distance from P1 to epicranial 
suture about % that from P1-L; L cephalad of juncture of adfrontal ecdysial lines. Ocellar 
spacing: Ocl—Oc2 0.053-0.058 mm; Oc2-—Oc3 0.034-0.039 mm; Oc3-—Oc4 0.029-0.034 
mm. 

Mouthparts. Identical to those of H. uniformis. Hypopharyngeal complex (Fig. 4): 
spinneret very thin, transparent, tapering distally, about 1.5 times the length of Lpsl; 
Lp2 and about % the length of Lps1; stipular setae slightly more than % the length of 
Lpsl, slightly less than twice the length of Lpl, about equal to Lp2; Lps2 about % the 
length of Lp]; distal and proximal regions of hypopharynx continuous, no medial trans- 
verse cleft present; distal and proximolateral regions of hypopharynx covered with small 
fine spines, spines becoming slightly longer proximally. Mandible (Figs. 5, 6): two well- 
separated outer setae present; inner surface with distinct ridges; lacking inner tooth; with 
six outer teeth, the sixth rounded and indistinct, the first five angular and well developed; 
outer margins of all teeth lacking serrations. 

Thorax. Segment Tl (Fig. 9): prothoracic shield heavily sclerotized; SD1 and SD2 
setal insertions separated from the edge of the prothoracic shield; interspace D1-D1 
about 0.82 XD1-SD1; D2-SD2 about 1.57 SD2-XD2; seta L2 present, much finer than 
L1; spiracle elliptical, 0.19-0.20 mm high, 0.112-0.117 mm wide; peritreme wider lat- 
erally. T2 (Fig. 5): D1—D2 about 0.82 D2-SD2; all setae thin and hairlike, tapering and 
sharply pointed distally; coxal bases widely separated. T3: Ts3 spatulate (Fig. 9), not 
tapering distally as in H. uniformis. 

Abdomen. Dorsal and lateral chaetotaxy of Abl-10 as in Fig. 9. Abl with 2 SV setae, 
Ab2-6 with 3 SV setae, Ab7-8 with 1. Ab9: SD1 much finer and hairlike than D1 and 
D2. Ab10: anal shield as in Fig. 8. Dorsal margin convex, posterior margin entire. Length 
of D1 on Ab6-7 0.240-0.245 mm; D2 0.26-0.27 mm. Asp 7 0.12-0.13 mm high, 0.09 
mm wide; Asp8 0.325-0.328 mm high, 0.15 mm wide. 


308 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY 


OC 


Le WE 
0G 


ae 


als l Cie ZALES 

aT ee cece }s Bas 
Ao Sot | os \; | o/ 6 { ON ee 

474 ee Cee im 
pe a OQ 
AG ieniasS e\5 Za 

9 T2 Ab6 Ab7 Ab8 Ab9 
Ab10 


Fics. 8-9. Homorthodes furfurata, larval structures: 3, head capsule, frontal view; 
4, hypopharyngeal complex, left lateral view; 5, left mandible, oral surface; 6, left 
mandible, outer surface; 7, left mesothoracic tibia and tarsus; 8, anal shield, dorsal view; 
9, dorsolateral chaetotaxy of prothoracic (T1), mesothoracic (T2), and abdominal seg- 
ments (Ab1—2, Ab6-10). Scale lines equal 1.0 mm. 


Material examined. 2 specimens: 2.5 km south of Tomahawk Lake, Halifax Co., Nova 
Scotia. Reared on Taraxacum officinale Weber from ova obtained from a female on 29 
July 1978. Larvae pupated 8-10 October 1978. Moth collected, determined, and larvae 
reared by K. A. Neil. 

Remarks. The larvae of H. furfurata and H. uniformis are very similar and based 


VOLUME 37, NUMBER 4 309 


on the figures of H. uniformis given by Godfrey (1972), cannot be separated using head 
capsule and mouthpart structures. The spatulate Ts3 (Fig. 9) can be used to differentiate 
H. furfurata from H. uniformis, the latter having simple hairlike tarsal setae. Homor- 
thodes lindseyi (Benjamin) has Ts3 spatulate, but can easily be separated from H. fur- 
furata by the shorter tarsal setae, mandibular, and hypopharyngeal complex differences. 


ACKNOWLEDGMENTS 


I would like to thank Dr. G. L. Godfrey of the Illinois Natural History Survey for 
reviewing this manuscript, and Ronald Long of Simon Fraser University for photography. 


LITERATURE CITED 


CRUMB, S. E. 1956. The larvae of the Phalaenidae. U.S. Dept. Agr. Tech. Bull. 11385. 
356 pp. 

FERGUSON, D. C. 1954. The Lepidoptera of Nova Scotia: (Macrolepidoptera). Nova 
Scotia Mus. Sci. Bull. 2. 214 pp. 

FORBES, W. T. M. 1954. Lepidoptera of New York and neighboring states. Pt. III. 
Cornell Univ. Agr. Expt. Sta. Mem. 329. 433 pp. 

GopFREY, G. L. 1972. A review and reclassification of larvae of the subfamily Had- 
eninae (Lepidoptera, Noctuidae) of America north of Mexico. U.S. Dept. Agr. Tech. 
Bull. 1450. 265 pp. 

GROTE, A. R. 1874. New species of North American Noctuidae. Proc. Acad. Nat. Sci. 
Phil. 26:197-214. 

Hinks, C. F. & J. R. BYERS. 1976. Biosystematics of the genus Euxoa (Lepidoptera: 
Noctuidae). V. Rearing procedures and life cycles of 36 species. Can. Entomol. 108: 
1345-1357. 

ROCKBURNE, E. W. & J. D. LAFONTAINE. 1976. The cutworm moths of Ontario and 
Quebec. Can. Dept. Agr. Publ. 1593. 164 pp. 


Journal of the Lepidopterists’ Society 
37(4), 1983, 310-311 


GENERAL NOTES 


OBSERVATIONS OF HILLTOPPING MITOURA SPINETORUM AND 
M. JOHNSONI (LYCAENIDAE) IN CALIFORNIA 


Mitoura spinetorum (Hewitson) and Mitoura johnsoni (Skinner) are medium-sized 
hairstreak butterflies having broad distributions but low-density and sometimes localized 
populations. These closely related species have similar reproductive biologies, and both 
use hilltopping behavior as a mate-locating strategy. Past observations of hilltopping 
behavior by these species are summarized, and new observations of some California 
populations are presented. 

Previous reports of M. spinetorum behavior include hilltopping by males in the Prov- 
idence Mountains, San Bernardino County, California, and Black Ridge, Mesa County, 
Colorado (Shields, 1965, J. Res. Lepid. 4:233-250); hilltopping by both sexes in the eastern 
Mojave ranges (Emmel & Emmel, 1973, Butterflies of Southern California, Nat. Hist. 
Museum Los Angeles Co., Science Series 26:1-148); perching on small pines near hilltops 
in Grand Canyon, Arizona (Scott, 1973, J. Lepid. Soc. 27:283-287); and perching by 
males on ridgetop pinyons and junipers in Nevada (Austin & Austin, 1981, J. Res. Lepid. 
19:1-63). Shields (in Scott, 1973) observed M. johnsoni males perching on the tops of 
tall trees on a hilltop next to Thompson Canyon, Yolo County, California. 

Our observations of hilltopping adults were made at two locations in the Inner Coast 
ranges of California: North Peak (1080 m) of Mt. Diablo in Contra Costa County and a 
ridge (310 m) at Butts Canyon in Napa County. Digger pine (Pinus sabiniana Douglas) 
and California juniper (Juniperus californica Carriere) occur in a semi-open area on the 
summit of North Peak. Only one conifer, digger pine, grows on the ridgetop at Butts 
Canyon in serpentine chaparral. Digger pine is commonly parasitized by pine dwarf 
mistletoe (Arceuthobium campylopodum Engelmann), the larval host plant of M. spi- 
netorum and M. johnsoni in the study areas. Mitoura spinetorum occurs at both North 
Peak and Butts Canyon; however, M. johnsoni has been found only at the latter location. 

Mitoura spinetorum appears to have three broods on North Peak. Adults have been 
found at this location from late March to early May, June to early July, and mid-August 
to late September (Opler & Langston, 1968, J. Lepid. Soc. 22:89-107; our observations 
1975-1981). At Butts Canyon, adult M. spinetorum and M. johnsoni have been found 
in March, April, and June (Langston, pers. comm.; our observations, 1981). Although no 
late-season individuals of either species have been encountered at this site, it seems likely 
that at least M. spinetorum has a third emergence at Butts Canyon. 

On North Peak, M. spinetorum males perch on the needles and occasionally on sta- 
minate cones of digger pine and foliage of junipers growing about the summit and nearby 
ridgetops. Some of these trees appear to be more often used for perching than others, as 
adults have consistently been found on the same conifers over several seasons. Trees used 
for perching are not necessarily the tallest present, may or may not have mistletoe, and 
are always located on a ridge crest but not necessarily at the highest point. At Butts 
Canyon, male M. johnsoni perch mostly about the tops of digger pines approximately 
six meters in height. Many of these trees are infested with pine dwarf mistletoe, some 
heavily. 

Mitoura spinetorum males perch by alighting for periods of up to 35 minutes during 
which they may wave the antennae, rub the hindwings together, shift body orientation, 
or remain motionless. Males fly from perching sites to investigate conspecific males and 
other insects (lycaenids, hesperiids, and dipterans), to circle erratically about the perching 
site, or to transfer to other perching sites. One M. spinetorum was continuously engaged 
in these activities from 0920 to 1105 hours PST (Sept.) on North Peak. Mitoura johnsoni 
males exhibit similar perching behavior. 


! Records of Clark County, Nevada M. spinetorum supplied by an anonymous reviewer also show a disproportionate 
sex ratio at hilltops (24 males, 0 females) and at canyon bottoms (4 males, 11 females). 


VOLUME 37, NUMBER 4 811 


Females were seen at our study sites only a few times. On 4 April 1981 two mating 
pairs of M. spinetorum were found on North Peak. Both were on trees frequently used 
by perching males. One pair was resting on juniper foliage, the other on a staminate 
cone of digger pine. Occasional North Peak females were seen flying about pine dwarf 
mistletoe on a ridgetop near the summit. Although no mating M. johnsoni were found 
at Butts Canyon, several ovipositing females were observed on the ridgetop in April. The 
disproportionate sex ratio at the ridgetops and summit may indicate female dispersal 
after mating.! 

These observations are consistent with Shields’ (1967, J. Res. Lepid. 6:69-178) and 
Scott’s (1970, J. Res. Lepid. 7:191-204) conclusions that butterflies with low population 
densities hilltop in order to facilitate the rendezvous of mates. 

We thank Maria F. Lane, Robert L. Langston, Larry J. Orsak, Jerry A. Powell, and 
Arthur M. Shapiro for their assistance. 


RICHARD V. KELSON, 29 Tiffin Ct., Clayton, California 94517 AND MARC C. MINNO, 
Aquatic Plant Management Laboratory, 3205 SW 70th Avenue, Fort Lauderdale, Flor- 
ida 33314. 


Journal of the Lepidopterists’ Society 
87(4), 1983, 311-313 


LEPIDOPTERA REARED ON A SIMPLE WHEAT GERM DIET 


Artificial diets have been used as food in rearing many species of Lepidoptera (Singh, 
1972, Bull. N.Z. Dept. Scient. Ind. Res. 209 pp.; Vanderzant, 1974, A. Rev. Entomol. 19: 
139-160; Hinks & Byers, 1976, Can. Entomol. 108:1345-1357). They may be synthetic 
(meridic), or composed of one or more natural products (oligidic) such as wheat germ 
and homogenized beans. The latter type is especially useful in rearing the larvae of 
polyphagous species of Lepidoptera, since no specific phagostimulants are required. 

Fifty-seven species of Lepidoptera, mainly Noctuidae but also Lymantridae and Geo- 
metridae (Table 1), were reared from egg to adult on a simple wheat germ diet from 
1977-1980. Adult females were collected at either a 15 watt ultraviolet light or sugar 
bait. Females thus collected were placed in 10 x 6 x 2 cm clear polystyrene boxes and 
fed a 10-15% sucrose solution until eggs were laid. 

Larvae of all species were fed an artificial diet based on that of Hinks and Byers 
(1976), except that kidney beans were used instead of pea beans. An additional 100 g of 
wheat germ and 12 ml of formaldehyde were also used. The formaldehyde had no effect 
on the growth of any species of Lepidoptera bred, although it is known to have an 
inhibitory effect on the growth of other kinds of insects (Singh & House, 1970, J. Insect 
Physiol. 16:1969-1982). 

Rearing techniques followed those developed by Hinks and Byers (1976) for the genus 
Euxoa, except that larvae were reared in 10 x 6 x 2 cm clear polystyrene boxes, with 
15-20 larvae/box. At the fourth instar the larvae were separated and reared to maturity 
individually in 15 x 100 mm disposable polystyrene Petri dishes. All larvae were reared 
at 25-30°C under a photoperiod of 15-9 h light-dark cycle. 

Feeding was discontinued at the first visible signs of the prepupal period, and 5- 
10 larvae were placed in 946 ml polystyrene containers partially filled with moist, ster- 
ilized top soil. A strip of paper towel provided a vertical surface for the moths to crawl 
up upon emergence. The containers were sealed with clear polyethylene and were kept 
at the same temperature and photoperiod conditions as the larvae. 

Newly eclosed larvae of two noctuid species, Feralia comstocki Grt., a general feeder 
on coniferous trees, and Homorthodes furfurata (Grt.) which has been recorded from 
Acer spp. (Rockburne & Lafontaine, 1976, The Cutworm Moths of Ontario and Quebec. 


312 


TABLE l. 


Family 


Geometridae 


Lymantriidae 


Noctuidae 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Species 


Melanolophia canadaria (WIlk.) 
Pero morrisonaria (Hy. Edw.) 


Orgyia leucostigma plagiata 
(W1k.) 


Orgyia leucostigma sablensis Neil 


Zale minerea (Gn.) 
Autographa flagellum (W1lk.) 
Plusia putnami Grt. 
Acronicta innotata Gn. 
Acronicta noctivaga Gn. 
Crymodes devastator (Brace) 
Phlagophora iris Gn. 
Elaphria festivoides (Gn.) 
Xylena curvimacula (Morr.) 
Lithophane innominata (Sm.) 
Eupsilia vinulenta (Grt.) 
Eupsilia tristigmata (Grt.) 
Eupsilia morrisoni (Grt.) 
Siderides maryx (Gn.) 

Polia imbrifera (Gn.) 

Polia latex (Gn.) 

Melanchra adjuncta (Gn.) 
Melanchra assimilis (Morr.) 
Lacanobia atlantica (Grt.) 
Lacanobia grandis (Gn.) 
Lacanobia lutra (Gn.) 
Lacanobia legitima (Grt.) 


Lacanobia lilacina (Harv.) 


Lacinipolia renigera (Steph. ) 
Lacinipolia lorea (Gn.) 
Lacinipolia olivacea (Morr.) 
Pseudaletia unipuncta (Haw.) 
Leucania inermis (Fbs.) 
Crocigrapha normani (Gtrt.) 
Orthosia revicta (Morr.) 
Orthosia hibisci (Grt.) 
Homorthodes furfurata (Grt.) 
Pseudorthodes vecors (Gn.) 
Tricholita signata (Wlk.) 


Agrotis volubilis Harv. 


Agrotis ipsilon (Hufn.) 
Feltia heralis (Grt.) 
Euxoa divergens (Wlk.) 
Euxoa scandens (Riley) 
Euxoa tristicula (Morr.) 
Euxoa perpolita (Morr.) 
Ochropleura plecta (L.) 


Lepidoptera reared on artificial wheat germ diet. 


Larval foodplant 


General feeder 
General feeder 


General feeder 


General feeder 


General feeder on deciduous trees 
Helianthis, Liatris: 

Grasses 

General feeder on deciduous trees 
General feeder on deciduous trees 
Grasses 

General feeder 

General feeder on deciduous trees 
General feeder 

General feeder or deciduous trees 
General feeder 

General feeder 

General feeder 


General feeder on deciduous trees 

General feeder on deciduous trees 

General feeder 

General feeder 

General feeder on low plants 

General feeder 

General feeder 

Grasses, general feeder on low 
plants 

Grasses, general feeder on low 
plants 

General feeder 

General feeder 

General feeder on low plants 

Grasses, general feeder 

Grasses 

General feeder on deciduous trees 

General feeder on deciduous trees 

General feeder on deciduous trees 

Acer spp.’ 

General feeder on low plants 

Feeds in stems and flowers of 
various Compositae 

Achillea millefolium L., 

Vaccinium vacillans Torr. 

Oenothera biennis L.4 

General feeder 

General feeder 


General feeder 


Genera! feeder on low plants 


VOLUME 37, NUMBER 4 3138 


TABLE 1. Continued. 


Family Species Larval foodplant 
Peridroma saucia (Hbn.) General feeder 
Diarsia jucunda (Wlk.) Grasses 
Eurois astricta Morr. General feeder on woody plants 
Xestia dolosa Franc. General feeder 
Xestia normaniana (Grt.) General feeder 
Xestia smithii (Snell.) General feeder 
Xestia oblata (Grt.) General feeder on low plants 
Metalepsis fishii (Grt.) Vaccinium?2* 
Anaplectoides prasina (D. & S.) General feeder on low plants 
Eueretagrotis perattenta (Grt.) General feeder 


' Tietz, 1972, An Index to the Described Life Histories, Early Stages and Hosts of the Macrolepidoptera of the 
Continental United States & Canada. 11, A. C. Allyn, Sarasota, 1041 pp. 

2 Rockburne & Lafontaine (1976). 

3 Ferguson, 1975, U.S. Dept. Agric. Tech. Bull. 1521, 49 pp. 

4McCabe, 1981, J.N.Y. Entomol. Soc. 89(2): 59-64. 


Can. Dept. Agric. Publ. 1598. 164 pp., 613 figs.) refused to eat the artificial diet. Both 
species were subsequently reared on previously recorded host plants. 


KENNETH NEIL, Department of Biological Sciences, Simon Fraser University, Bur- 
naby, British Columbia, CANADA V5A 1S6. 


Journal of the Lepidopterists’ Society 
87(4), 1983, 313-317 


EGG PLACEMENT BY PHOEBIS (PIERIDAE) ON CASSIA (LEGUMINOSAE) 
“ANTICIPATES” THE TROPICAL RAINY SEASON 


As the tropical dry season advances, one common response by green plants is a pro- 
gressive loss of leaves, which is sometimes accompanied by a gradual development of 
new leaf buds near the end of this period (Janzen, 1967, Evolution 21:620-637). By the 
time the rainy season is underway, such a plant species exhibits considerable leafing-out, 
providing herbivorous insects with an expanded food base. At any given Central Amer- 
ican locality, not all plant species respond to the same degree to the dry season. 

At “Finca La Tigra” on the Atlantic slope (220 m elev.) of Costa Rica’s Cordillera 
Central (near La Virgen, Heredia Province, 10°23’N, 84°07'W) the plant family Legu- 
minosae exhibits a broad range of dry season response patterns during the longer of two 
dry periods characteristic of this Premontane Tropical Wet Forest locality (Fig. 1). Many 
legume genera remain fully leaved throughout the major dry season, although the pro- 
duction of new leaf buds is often low. Others remain evergreen and exhibit considerable 
leaf replacement at this time. Still others, such as the roadside shrub or small tree (canopy 
height 2-5 m), Cassia fructicosa Mill., exhibit considerable loss of mature leaves, followed 
by a gradual development of new leaf buds in the latter part of the major dry period, 
which usually extends from late December through March. This is also a period of greatly 
reduced flowering and complete absence of fruits on C. fructicosa (Allen M. Young, 
unpubl. data, 1973-1982). The guild or assemblage of herbivorous insects associated with 
this tree species must cope physiologically and behaviorally with this annual cycle of 
seasonal changes in the availability of various edible plant parts. 


314 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


450 


400 


350 


300 


250 


Mean Monthly Rainfall (mm) 


J F M A M J J A S O N D 
Months 


Fic. 1. Pattern of monthly rainfall at “Finca La Tirimbina,” near La Virgen, Heredia 
Province, Costa Rica. Note the short dry season occurring in February and March. 


In this note, I report that the pierid butterfly Phoebis argante (Fabricius), which is 
very abundant at this locality as it is throughout much of Costa Rica and Central America 
overall, behaviorally circumvents the deciduous habit of C. fructicosa by preferentially 
placing its eggs singly on the developing leaf buds while avoiding the severely thinned- 
out canopy of mature leaves. On one roadside stretch at La Tigra, I observed frequent 
egg placement on C. fructicosa by P. argante on roughly 30 trees between 1978 and 
1982, providing a sample of 68 observed oviposition acts in the major dry season (De- 
cember—March). In addition, 35 oviposition acts were observed during the lengthy rainy 
season characteristic of this region (Fig. 1). 

During the rainy season, P. argante frequently places eggs singly on the fresh, fully- 
expanded leaves of C. fructicosa, as it does for other larval food plants at this locality 
during both rainy and dry seasons (A.M.Y., unpubl. data). A good example of the latter 
is Pentaclethra macroloba, a legume tree that remains evergreen throughout the dry 
seasons at this locality. But in the case of C. fructicosa as a larval food plant, during the 
latter part of the major dry season eggs are placed singly only on the small (length 4-10 
mm, n = 30 measured on 7 March 1982) leaf buds that are scattered below the canopy 
of old leaves. A female butterfly weaves through the branches of the tree until it finds a 
new leaf bud and carefully places an egg on it (Fig. 2). A butterfly may thus deposit 
anywhere from 1 to 6 eggs on a single tree during one visit. Leaf buds having eggs are 
sometimes avoided, although a single female may place multiple eggs on a given bud. 
Butterflies do not even “inspect” the older leaves of a tree’s canopy (Fig. 2). 

The stage at which P. argante places eggs on leaf buds is followed by a period of 


VOLUME 387, NUMBER 4 


. "Eor 
yp«-€¢ e a Je * : ~ 


Fic. 2. Above: Position of eggs of Phoebis argante on unfolding new leaf buds of a 
larval host plant, Cassia fructicosa (Leguminosae) at “Finca La Tigra” in March. Below: 
The thinned-out canopy of mature leaves of Cassia fructicosa as it appears during 
February and March at “Finca La Tigra.” 


316 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


rapid growth and expansion of the buds, a condition that precludes the necessity of a 
potential egg diapause at this locality. The degree to which Neotropical pierids have 
diapausing eggs is unknown. The condition of rapid leafing-out is analogous to the avail- 
ability of new leaf buds precluding the need for a reproductive diapause mechanism in 
the adults of butterfly species at such a locality. Eggs deposited on leaf buds in late 
February generally hatch in less than 10 days, and by this time the new leaves are 
considerably larger (expanded). Fresh adults of P. argante are found at this locality 
throughout the year, indicating a continuous breeding population structure made possible 
by (1) the behavioral flexibility that allows P. argante to place eggs on several alternate 
leguminous food plants at any one time, and (2) behavioral flexibility permitting different 
types of egg placement on a food plant that exhibits a markedly deciduous habit in at 
least the major dry season. Placing eggs on new leaf buds juxtaposes newly hatched larvae 
with a fresh food supply, and perhaps does so before other members of the herbivore 
guild associated with the leaves of C. fructicosa discover this resource at the beginning 
of the rainy season. 

Such observations suggest that flexibility in egg placement behavior in a butterfly 
species that exploits a food plant with a deciduous habit is an effective mechanism by 
which the insect “anticipates” the expanded food supply of fresh, and perhaps, more 
edible, leaves, that will be available at the beginning of the tropical rainy season. Such 
a behavioral response permits the butterfly to breed throughout the year at such a locality. 
As with many Neotropical butterflies, P. argante possesses an egg-to-adult developmental 
period of about 40 days, including a larval period of 22 days, a period sufficiently long 
enough to “pace” development with the growth of new, highly edible, leaves of the 
larval food plant. Because this butterfly is geographically widespread throughout the 
Neotropical Region (Seitz, 1924, Macrolepidoptera of the World, Vol. 5: American Rho- 
palocera, Stuttgart: A. Kernan, 615 pp.; Howe, 1975, The Butterflies of North America, 
New York: Doubleday, 633 pp.), and because it exploits a range of Cassia species (d’Al- 
meida, 1940, Arq. de Zool. Estado de Sao Paulo, Revista do Museu Paulista 1:67-152; 
Biezanko, 1959, Arq. de Entomol. Minist. Agric. Brasil. Ser. B; Teitz, 1972, An Index to 
the Described Life Histories, Early Stages and Hosts of the Macrolepidoptera of the 
Continental United States and Canada, Allyn Museum, 1041 pp.; Howe, op. cit.), regional 
or local larval food plant differences and the degree of seasonality are major determinants 
of whether or not the insect will exhibit the form of egg placement behavior described 
in this note. It is well known that many Lepidoptera species preferentially place eggs on 
fresh leaves of the food plant, regardless of a seasonal cycle in the availability of new 
leaves and other vegetative parts used as food. Presumably such plant parts are more 
edible for small caterpillars in the sense of having fewer defensive compounds (concen- 
tration, or range in types) and other barriers to successful feeding. What this present 
paper suggests is that in tropical regions with one or more dry seasons, we must view 
such preferences through the evolutionary lens of effects of the dry period on the supply 
of fresh leaves. 

Not all legume-feeding Lepidoptera at this locality preferentially oviposit on young 
or fresh leaves. Even though the common woody legume vine Machaerium seemannii 
also flushes out new leaves in the dry season here, Morpho peleides limpida Butler and 
Morpho granadensis polybaptus Butler (Morphidae) place their eggs singly on the ma- 
ture leaves of the vines (Young & Muyshondt, 1978, Carib. J. Sci. 13:1-49; Young, 1974, 
J. Lepid. Soc. 28:90-99; Young, 1982, J. New York Entomol. Soc. 90:35-54), even in the 
presence of young leaves. Newly flushed leaves in a leguminous food plant, therefore, 
may not always be more biologically suitable as larval food than the mature leaves of 
the same plant. But in the case of the P. argante x C. fructicosa association in Central 
America, newly flushed leaves may be more suitable as larval food than older leaves on 
the same trees. When 10 newly hatched first-instar larvae of P. argante were offered the 
old leaves of C. fructicosa that were present on trees in late February, some larvae 
nibbled at the edges of leaves, but all died within two days. A simultaneously studied set 
of another 10 first-instar larvae reared on the leaf buds where eggs were placed by 
ovipositing butterflies all developed normally during the same period, and none died. 
Clearly in this case, if we eliminate the bias of small sample size, older leaves of C. 
fructicosa available late in the dry season in Costa Rica are markedly unsuitable for 


VOLUME 37, NUMBER 4 SZ 


proper larval development in P. argante than newly unfolding leaves available at the 
same time. I suspect that full-sized mature leaves of C. fructicosa available in the rainy 
season are also highly unsuitable to P. argante larvae, before these leaves assume the 
brittle and blotched appearance that characterizes them in the dry season. 


ALLEN M. YOUNG, Invertebrate Zoology Section, Milwaukee Public Museum, Mil- 
waukee, Wisconsin 53288. 


Journal of the Lepidopterists’ Society 
87(4), 1983, 317-318 


CHERMOCK, HOVANITZ AND WEBER COLLECTIONS 
DONATED TO ALLYN MUSEUM 


Among other accessions during 1980 and 1981 received by the Allyn Museum of 
Entomology of the Florida State Museum were three large and very significant collections 
of Rhopalocera donated by the heirs of Franklin H. Chermock, William Hovanitz and 
Bernard H. Weber. All of these collections have filled great gaps in the coverage of the 
Museum's holdings, especially in Arctic species and in the specialty groups of each of 
the donors. 

Franklin H. Chermock collected and studied Lepidoptera assiduously and enthusias- 
tically for over forty years until his death in 1967. Much of his early taxonomic work 
was done in collaboration with his brother, the late Ralph L. Chermock, and their 
investigations into the fauna of the Riding Mountains, Manitoba, were enormously valu- 
able scientifically, resulting in the descriptions of many endemic butterflies from that 
region. Dr. Chermock’s interest in Canadian butterflies continued up to the time of his 
death, and in company with his son, the late Paul W. Chermock, Frank collected and 
studied the butterflies throughout northern Manitoba and much of the Northwest Ter- 
ritories. The Chermocks, father and son, intended to describe many new taxa from these 
expeditions and distributed innumerable manuscript “paratypes of these butterflies, 
many of which have been named subsequent to Frank’s death by other authors. The 
Chermock collection contained 56,154 specimens (nearly all from the Nearctic), includ- 
ing 20 holotypes, eight allotypes, five syntypes and nearly 2000 paratypes, many from 
other authors. The “type series” of 56 proposed taxa were included and labelled. About 
1300 microscopic genitalia slides and a useful library augmented the collection itself, 
along with a sizable body of correspondence relating to it. The Chermock material has 
provided the Allyn Museum collection with its first significant holdings in Arctic and 
Subarctic butterflies, and we are grateful to Frank’s daughter, Mrs. Linda C. Hassinger, 
for the opportunity to preserve it and make it available for study. 

William Hovanitz collected and studied Rhopalocera for about forty years before his 
untimely death in 1977. He was best known for his genetic and variational studies; the 
personally collected material for these studies is preserved in his collection. He had no 
parochial bias, and those groups that were of special interest to him are represented in 
the collection from throughout their ranges, especially Colias, Argynninae and Oeneis. 
The Hovanitz material included 23,859 specimens, including more than 4400 Colias, 
significant numbers of which were from the Arctic of Canada and Alaska and from 
outside the Nearctic (especially the Andes of Peru, Bolivia, Chile and Argentina). One 
of the most valuable parts of the collection is material taken along a transect of the Alaska 
Highway from about Grande Prairie, Alberta (before the beginning of the Highway) to 
Tok Junction, Alaska. While this collection is particularly strong in Colias, other groups 
are well represented, such as Clossiana, Oeneis and the “blues.” The only type material 
in the collection were two paratypes of Colias thula Hovanitz and two Bang-Haas Colias 


318 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


cotypes. Our gratitude to Dr. Hovanitz’ widow, Barbara, for this generous gift is here 
acknowledged. 

Bernard H. (Bernie) Weber was an avid collector who also carried on an ambitious 
exchange program with other lepidopterists, chiefly throughout the United States and 
Canada. His collection is composed of 43,988 specimens, more than 90% of which are 
Nearctic and represent the majority of the presently recognized taxa of the region. Many 
so-called “rare” butterflies (such as Speyeria egleis tehachapina J. A. Comstock) are 
included in the collection in sizable series. California and West Coast butterflies are 
especially well represented, but there are good series from elsewhere as well; no type 
specimens are included. This collection came to the Museum through the kindness and 
foresight of Mr. Weber's widow, Virginia. 

All of these collections have been or are being incorporated into the main collection 
of the Allyn Museum of Entomology where they will be available to researchers in the 
coming years. Suffice it to say that the staff of the Museum is grateful for the generosity 
(that human attribute that makes growth of museum collections possible) and far-sight- 
edness of the donors in making these collections again available to Science. 


LEE D. MILLER, Curator, Allyn Museum of Entomology, Florida State Museum, 
3701 Bay Shore Road, Sarasota, Florida 33580. 


Journal of the Lepidopterists’ Society 
37(4), 1983, 318 


AN OLD FIRST UNITED STATES RECORD FINALLY PUBLISHED: 
PAPILIO VICTORINUS (PAPILIONIDAE) IN LAREDO, TEXAS 


On 17 August 1974 I collected a male Papilio victorinus Dbldy. in Laredo, Webb 
County, Texas. The relatively fresh specimen was collected in a large planted flower bed, 
which included a number of marigolds on the eastern edge of the city. 

I first mentioned this capture in the May/June 1978 issue of the “News of the Lepi- 
dopterists’ Society.”’ At that time I was inquiring as to whether anyone else had collected 
this particular species in the U.S. I did not receive any correspondence as to such captures; 
subsequently, the publication of the 1981, Catalogue/Checklist of the Butterflies of 
America North of Mexico by Miller & Brown, with the omission of P. victorinus, recon- 
firmed my belief that this was indeed a first U.S. record. 

Tyler (1975, The Swallowtail Butterflies of North America, Nature Graph Publishers, 
Inc., Healdsburg, California, p. 128) gives the range of this species as “Temperate and 
tropical regions of E. Mexico as far north as Nuevo Leon and Tamaulipas.’ Five addi- 
tional specimens were collected during a short period in August 1974 on the Plaza 
Zaragosa in Monterrey, Nuevo Leon, Mexico. I had expressed the belief that P. victorinus 
was perhaps extending its range northward; however, more recent trips to northern 
Mexico have failed to turn up the species in any significant numbers. It appears that 
1974 was just a good year for it. The fact that it can be found occasionally in good 
numbers in Monterrey, which is just south of the latitude of Brownsville, Texas, is an 
indication that collectors in southern Texas should watch for it. 

The specimen is presently in my collection at 185 N. Missouri St., Liberty, Missouri 
64068. 


JAMES K. ADams, Dept. of Entomology, University of Kansas, Lawrence, Kansas 
66045. 


VOLUME 37, NUMBER 4 319 


Journal of the Lepidopterists’ Society 
37(4), 1988, 319-320 


ESTIMATION OF DAILY OVIPOSITION RATES 
IN REARED FEMALE ANTHERAEA POLYPHEMUS (SATURNIIDAE) 


In rearing giant silkworm moths for research purposes, it is frequently important to 
use specimens having the same approximate age and physiological condition. Funda- 
mental to obtaining such specimens is an understanding of oviposition patterns and the 
factors that may affect such patterns. We have studied oviposition in Antheraea poly- 
phemus (Cramer) (Miller & Cooper, 1980, J. Lepid. Soc., 34:256-259) and determined 
that reared females live an average of six days after mating and deposit an average of 
216 eggs. Peak oviposition is during the first three nights after mating. We have also 
determined (Miller, et al., 1982, Ann. Entomol. Soc. Amer., 75:107-108) that the number 
of mature eggs (NME) in reared females can be estimated from pupal weight (WT) by 
the linear regression equation: NME = 2.22 + 45.9 WT (r = 0.78). The number of 
mature eggs at emergence is 74.0 percent of the total eggs (TE) and is related by the 
linear regression equation: TE = 35.3 + 1.10 NME (r = 0.96). Studies have now been 
completed on two additional aspects of egg production in A. polyphemus: (1) the rela- 
tionship between total eggs deposited and adult longevity and (2) the relationship be- 
tween total eggs deposited and daily oviposition rates. These studies were undertaken 
because it was important to know whether females that lived longer, or contained larger 
numbers of eggs at emergence, deposited their eggs over a longer period of time and, 


125 


= 6.44 + 0.27E 


Fa3 


T 


HOO R =0.94 


= 
w” 75 
> 
<x 
[@) 
fo 
Ww 
a 
50 
” 
i.e) 
(6) 
Ww 
25 


O 100 200 300 400 


TOTAL EGGS DEPOSITED (E-_) 


320 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


thus, for any particular group of eggs, resulted in wider variation in physiologic ages of 
the embryos and subsequent stages. Eggs were collected from 40 reared female A. poly- 
phemus in paper bags as described by Miller & Cooper (1980). Daily oviposition rates 
represent the average eggs per night for the first three nights after mating. Total eggs 
represent the total eggs deposited during the lifespan. 

Antheraea polyphemus females live from five to 10 days after mating (mean = 7.1; 
S.D. = 1.8). No significant correlation (r = 0.14) was found between the total number of 
eggs deposited and female longevity. However, daily oviposition rates were highly cor- 
related (r = 0.94) with the total number of eggs deposited (Fig. 1). The total number of 
eggs deposited averaged 231.6 + 67.7. The number of eggs deposited per night during 
the first three nights after mating averaged 68.5 + 19.4. Thus, females that deposited 
greater total numbers of eggs during their lifespan did not live longer and deposit them 
over a longer period of time. They deposited their eggs at a greater rate during the first 
three nights after mating. The relationship between the three-day average oviposition 
(E,,) and the total number of eggs deposited (E,) is described by the linear regression 
equation: E,, = 6.44 + 0.27 E,. Using the relationship we previously demonstrated (Miller, 
et al., 1982) between NME and WT, and between NME and TE, it is possible to estimate 
daily oviposition (E,,) on the basis of pupal weight (WT) as follows: 


Eas = 6.44 + (0.27 (22.4 + 45.9 WT)) 


Aside from the value of this information in rearing giant silkworm moths for research 
purposes, the adaptive significance of these findings must also be noted. It appears that 
A. polyphemus, that does not feed as an adult, is able to efficiently use stored energy 
reserves obtained in the larval stage to deposit the majority of its eggs (74 percent) in a 
relatively short period of time (three days) independent of the total number of eggs 
deposited or the life-span of the moth. 


THoMas A. MILLER, U.S. Army Medical Bioengineering R&D Laboratory, Fort De- 
trick, Maryland 21701 and William J. Cooper, Florida International University, Miami, 
Florida 33199 (the opinions contained herein are those of the authors and should not be 
construed as official or reflecting the views of the Department of the Army). 


Journal of the Lepidopterists’ Society 
37(4), 1983, 320-321 


ADDITIONAL COMMENTS ON THE BUTTERFLIES OF THE 
AUSTIN, TEXAS, REGION 


Recently, Durden (1982, J. Lepid. Soc. 36:1-17) presented and analyzed a list of 173 
species of butterflies and skippers from a ten county region centered around Austin, 
Texas. Special attention was given to the 128 species found in Barton Creek canyon in 
the Balcones Fault Zone area of Austin. Below are comments on two species attributed 
to this author’s collecting activities plus the report of an additional species to the Austin 
region. 

In his list, Durden (op. cit.) credited an Austin specimen of Siproeta (Victorina) 
stelenes biplagiata (Fruhstorfer, 1907) to “R. Neck.” Although stray individuals of this 
species undoubtedly occur in the Austin area on rare occasions, I have never collected 
biplagiata in Austin. The closest personal record of biplagiata to Austin is a female 
collected in Garner State Park, Uvalde County, Texas, on 14 October 1976. Collection 
was made in the canyon of the Frio River, approximately 210 kilometers from Austin. 
The specimen was extremely worn and exhibited very weak flight behavior. Both these 


VOLUME 37, NUMBER 4 Sol 


observations indicate that the specimen was a long distance dispersant organism, one of 
many individuals of several species of tropical affinity which move northward in late 
summer and autumn of years in which moisture is sufficient (Neck, 1978, J. Lepid. Soc. 
32:111-115). Any Austin specimens would be of similar origin. Note should be made of 
the report of this species by Parks (Engelhardt, 1934, Brooklyn Entomol. Soc. 29:16) at 
San Antonio (125 km to the south) following a major hurricane in south Texas (see Neck, 
1977, J. Lepid. Soc. 31:67-68). 

The report of Eueides isabellae zorcaon (Reakirt, 1866) credited to “R. Neck” refers 
to a specimen sighted (but not collected) on 19 August 1971 on the floodplain of the 
Colorado River next to presentday Town Lake. This tract of land was highly disturbed 
by human activities in 1971 (now occupied by Austin High School). Weedy plants with 
significant nectar resources inhabited the area and attracted numerous species of but- 
terflies. The most important nectar source was Verbesina encelioides, a plant which is a 
major nectar source for butterflies in central Texas (see Neck, 1977, J. Res. Lepid. 16: 
147-154). Fall 1971 was a time of prodigious northward movement by countless butter- 
flies of numerous species as a result of heavy rains in August following a period of extreme 
drought (Helfert, 1972, Entomol. News 82:49-52; Neck, ms. submitted to J. Lepid. Soc.). 
Kendall (1972, J. Lepid. Soc. 26:49-56) reported a number of records of zorcaon from 
southern Texas in 1968; northernmost specimen was from San Antonio (125 km south of 
Barton Creek area). Occurrence of these 1968 specimens was attributed to introduction 
and establishment of breeding populations due to environmental effects associated with 
Hurricane Beulah of September 1967. Impact of this hurricane on the butterfly fauna of 
the Austin area has been discussed (Neck, 1978, J. Lepid. Soc. 32:111-155). 

I do, however, have one species to add to Durden’s list of Austin area butterflies. I 
collected a moderately worn female Anteos chlorinde nivifera (Fruhstorter, 1907) on 10 
November 1970 within the confines of the Brackenridge Field Laboratory (BFL) of the 
University of Texas at Austin within the city limits of Austin. The BFL specimen was 
collected near the mouth of an unnamed creek whose lower reaches are flooded by Town 
Lake (Colorado River), only 3.5 km from the mouth of Barton Creek. A. c. nivifera is 
well-documented to be a long distance traveler and is known as far north as Colorado 
and Kansas (Brown, 1960; J. Lepid. Soc. 14:156; Field, 1938, Bull. Univ. Kansas Biol. 
Series 39(10):1—828). Parks (see Engelhardt, op. cit.) also reported this species from Austin 
following a major hurricane in southern Texas (Neck, 1977, J. Lepid. Soc. 31:67-68). A. 
c. nivifera was also found in Austin in 1968 (leg. W. P. Hard, specimen in R. O. Kendall 
coll.); this year followed a major hurricane (Beulah) in 1967 which caused alterations in 
the butterfly fauna of the Austin area (Neck, 1978, J. Lepid. Soc. 32:111-115; 1981, Ibid. 
35:22-26). 

Another comment is appropriate concerning the listing under Zerene cesonia (Stoll, 
1790) of form “stainkeae” Field. Durden was referring to the form with reduced mar- 
ginal melanic markings. This form is correctly referred to as form “immaculsecunda”’ 
Gunder and is periodically common especially in late summer and autumn when sub- 
stantial movement from Mexico occurs (see Neck, 1981, J. Lepid. Soc. 35:22-26). Form 
“stainkeae” is a “rather rare white female form . .. with change of color yellow to white” 
(Field, 1936, J. Entomol. & Zoology (Pomona College) 28(2):17-26), which has been 
illustrated by Kimball (1965, Lepidoptera of Florida, Fla. Dept. Agric. 363 pp. pl. 1, fig. 
22). Durden (pers. comm.) has collected these whitish forms in the Austin area. 

I thank R. O. Kendall for comments on this manuscript. 


RAYMOND W. NECK, Texas Parks and Wildlife Department, 4200 Smith School Road, 
Austin, Texas 78744. 


Journal of the Lepidopterists’ Society 
87(4), 1983, 322-3824 


OBITUARY 
LUCIEN HARRIS, JR. (1899-1983), A Tribute 


One of the country’s most dedicated naturalists in recent times, Lucien Harris, Jr., 
spent a lifetime in the study and conservation of the natural history of his native state, 
Georgia, and the Southeastern region, and inspired all who knew him by his wealth of 
knowledge and experiences, and his quiet manner coupled with a unique intensity of 
spirit that radiated from the man. 

Lucien Harris, Jr. was born on 9 September 1899 in Atlanta, Georgia to a long line of 
Georgians, his grandfather, Joel Chandler Harris (1848-1908), the southern folklore writ- 
er and author of the endearing Uncle Remus stories, being the most famous. Lucien 
grew up in Atlanta, graduated from Boys’ High School and subsequently attended the 
University of Georgia, in Athens. After brief military service during World War I, he 
married Louise Nichols in 1919 and settled permanently in Atlanta. Over the next three 
decades the Harrises traveled extensively throughout the United States. Although an 
outdoors person by nature, Lucien established his career in the publishing field and served 
as the Southeastern Manager for Macmillan Publishers for many years. In that capacity 
he was directly involved with the logistics of the publication of Gone With the Wind 
and became a lifelong friend of its author, Margaret Mitchell. However, throughout his 
adult life, very often accompanied by Louise and their two sons, Lucien Harris III and 
James Robin Harris, Lucien spent virtually every moment he could find on field trips, 
to collect and study the butterflies and moths of the state and region. 

Lucien’s fascination with nature and wildlife no doubt was sparked by the picturesque 
stories of the animal characters in his grandfather's books, with which he was familiar 
from his early childhood. However, in his childhood and adolescence he became keenly 
interested in the observation of birds. During his teens, Lucien regularly went on bird 
walks with a family friend and a source of great inspiration, Wallace Rogers, who was 
a Methodist minister and also a nature photographer of considerable talent, with whom 
Lucien would later collaborate in co-authoring (along with Woolford B. Baker) a series 
of three volumes, Southern Nature Stories, directed at introducing children in the pri- 
mary grades to the world of nature. 

While observing the birds during those forays, Lucien also became interested in but- 
terflies, which were abundant throughout Georgia in those years when most of the state 
was still undeveloped. Before he was out of his teens he found himself building a serious 
collection and dedicating increasing amounts of his time to collecting specimens over a 
continuously widening area of the state. In his twenties he established friendships with 
lepidopterists and other naturalists in the state and in the region, notably Dr. P. William 
Fattig, an entomologist, and Woolford B. Baker, a botanist, both at Emory University, 
Dr. F. Strohecker, an entomologist at the University of Miami, and Fred Naumann of 
Forsyth, Georgia, an enthusiastic lepidopterist. In 1929 Lucien called an informal meet- 
ing at Emory University of his friends in various branches of science who shared an 
interest in natural history and conservation and founded what came to be the Georgia 
Society of Naturalists, and he served as its president for a number of years. The group 
met regularly, published bulletins and organized field trips to explore diverse areas of 
Georgia, the largest state east of the Mississippi. One of those areas was the Okefenokee 
Swamp, and it was through the untiring efforts of Lucien and the Georgia Society of 
Naturalists that the swamp was designated a National Wildlife Refuge. It was also through 
the efforts and negotiations of the Society and of Herbert Lee Stoddard, Sr., one of its 
members, that the Tall Timbers Research Station, near Tallahassee, Florida, was founded. 
Today, its main building houses the Herbert Lee Stoddard, Sr. bird collection, and the 
Lucien Harris, Jr. butterfly and moth collections, containing virtually all the specimens 
he collected between 1930 and 1970. 

As Lucien’s interest in Lepidoptera grew to a full-time preoccupation in his teens and 
twenties, he became keenly aware of the dearth of publications on the butterflies and 
moths of the region. Indeed, there had been only one major book on Georgia's butterflies, 
the two volume study, The Rarer Lepidopterous Insects of Georgia, published in London 


VOLUME 37, NUMBER 4 323 


“LUCIEN HARRIS, JR., 1899-1983 


in 1797 by Dr. James Edward Smith, and illustrated by Georgia’s pioneer naturalist, John 
Abbot (1751-1840). In a real sense Abbot, who collected and reared many of the butterfly 
species of Georgia and depicted them in their natural size and colors, with their cater- 
pillars, chrysalids and foodplants, became Lucien’s mentor, and for years his goal was to 
collect and document all the Georgia species painted by Abbot, even into the 1950's and 
1960's, when collecting with fellow lepidopterists John C. Symmes, Stanley S. Nicolay 
and Fred Naumann. In the process, as many new records were added to those of Abbot, 
and new information was amassed on life histories, foodplants, distribution, etc., Lucien 
undertook to update Abbot’s publication. In 1931, A List of the Butterflies of Georgia 
was published, being essentially an annotated checklist of the species taken in the state 
up to that time. A revised edition, edited by Austin H. Clark, was published in 1950 as 
The Butterflies of Georgia, Revised, containing substantial new material and a compre- 
hensive bibliography prepared by Mr. Clark. These two publications, along with exten- 
sive new records by Lucien and many other collectors in the 1950’s and 1960's, laid the 
groundwork for his culminating work, The Butterflies of Georgia, published in 1972 by 
University of Oklahoma Press. This volume, in the format of a field guide, covers and 
illustrates every species of butterfly known to occur in Georgia (NOTE: Four new species, 
Erora laeta, Mitoura hesseli, Euphyes dukesi and Urbanus dorantes, have been added 
to the state’s fauna since the book’s publication.). As many collectors have long since 
realized, however, the book has validity for virtually the entire Southeastern region, and 
it includes information on distribution, life history, foodplants, habitats and capture 
records for each species listed. 

In The Butterflies of Georgia is a detailed account of the discovery by Lucien, Lucien 


324 JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


III and J. P. Knudsen of what turned out to be a new species of giant skipper (Mega- 
thymidae), which was described and named Megathymus harrisi by H. A. Freeman in 
1955, in recognition of Lucien’s achievement. 

In addition to these accomplishments, Lucien was a charter member of the Lepidop- 
terists’ Society, a founding member of the Georgia Ornithological Society, an associate 
member of the American Association for the Advancement of Science, as well as a 
member of the Florida Audubon Society, the Tennessee Ornithological Society and the 
Georgia Academy of Science. 

Lucien Harris, Jr. died on February 22, 1983, in a Decatur, Georgia nursing home 
after a long illness. He is survived by his wife, Louise, his sons Lucien [JI and James 
Robin, and their respective children. In addition to the greater part of his collection of 
butterflies and moths contained in the Tall Timbers Research Station, his material col- 
lected up to 1930 is housed at the Emory University Museum, Atlanta and the balance 
at the Fernbank Science Center, Decatur, Georgia. 

It has been truly rewarding and inspiring to have known Lucien Harris, Jr. personally, 
to have experienced his contagious enthusiasm for the butterflies and other wildlife he 
knew so intimately, and to have received his friendship and comraderie. I am also 
deeply indebted to both Lucien Harris III and James Robin Harris for the information 
and photograph they provided me for the preparation of this tribute. 


IRVING L. FINKELSTEIN, 425 Springdale Drive N.E., Atlanta, Georgia 30305. 


Journal of the Lepidopterists’ Society 


37(4), 1983, 325-328 


INDEX TO VOLUME 37 


(New names in boldface) 


Abaeis nicippe, 272 
Achaea catella, 91 

A. faber, 91 

A. lienardi, 91 
Achytonix epipaschia, 134 
Acleris gloverana, 136 
Acronicta innotata, 312 

A. noctivaga, 312 
Adams, J. K., 318 
Agonopterix alstroemeriana, 38 
Agraulis vanillae incarnata, 272 
Agrotis arenarius, 14 

A. ipsilon, 312 

A. stigmosa, 14 

A. volubilis, 14, 312 
Agyrotaenia dorsalana, 137 

A. klotsi, 137 

A. provana, 137 
Albuna fraxini, 87 
Ambrosia trifida, 81 
Anacamptodes herse, 217 
Anaplectoides prasina, 312 
Antennaria aprica, 20 

A. neodioica, 20 
Anteos chlorinde, 89 

A. c. nivifera, 321 
Antheraea polyphemus, 319 
Anticarsia gemmatalis, 217 
Anua mejanesi, 91 

A. tirhaca, 91 
Apatura proserpina, 181 
Apolema carata, 146 
Argynnis apacheana, 79 
Argyrostagma niobe, 91 
Aristolochia californica, 237 

A. reticulata, 81 

A. serpentaria, 81 

A. tagala, 83 
Ascotis reciprocaria, 91 

A. selenaria, 91 
Asimina parviflora, 81 
Aspitates ochrearia, 146 
Asterocampa antonia, 270 

A. celtis, 269 

A. clyton, 145, 269 
Athletes ethra, 91 
Atlides halesus estesi, 272 
Austin, G. T., 244 
' Autographa flagellum, 312 
Barrows, E. M., 265 


Battus philenor, 81, 90, 236, 272, 275 


B. zetides, 171 
Beck, A. F., 89, 289 
Berenbaum, M., 38, 81 


Bitzer) hey jee 

Blanchard, A., 140 

Bombacopsis bipars, 91 
B. quinatum, 155 


Book Reviews: 28, 69, 95, 96, 187, 189, 259 


Bowe, J. J., 206 
Brachymeria ovata, 239 
Brown, F. M., 28, 37, 80, 181 


Brown, R. L., new hosts for Olethreutinae, 


224 

Bunaea alcinoe, 91 
Calhoun, J. V., 168 
Callaghan, C. J., 254 
Callophrys hesseli, 78, (97) 
Calycopis cecrops, 97 
Carmenta anthracipennis, 87 

C. haematica, 194 

C. mimosa, 194 

C. prosopis, 203 
Carolin, V. M., 129 
Cassia fructicosa, 318 
Celtis laevigata, 145 
Cercyonis oetus, 92 
Chionodes abella, 130 
Chlosyne gorgone carlota, 80 

C. lacinia adjutrix, 272 

C. nycteis, 85 
Choristoneura fumiferana, 169 
Chrysophanus dione, 181 
Clarke, J. F. G., 155, 224 
Cleora dargei, 91 

C. herbuloti, 91 

C. nigrisparsalis, 91 

C. rothkirchi, 91 

C. scobina, 91 
Clepsis persicana, 137 
Coleotechnites sp., 130 
Colias alexandra, 92, 178 

C. meadii, 177 

C. philodice eriphyle, 177 

C. thula, 317 
Colocleora divisaria, 91 
Colubrina texensis, 269 
Conium maculatum, 38 
Cooper, W. J., 319 
Crocigrapha normani, 312 
Crymodes devastator, 312 
Cypripedium calceolus, 265 

C. reginae, 265 
Danaus gilippus strigosus, 272 

D. plexippus, 92, 272, 275 
Dasychira basalis, 91 

D. georgiana, 91 
Dornfeld, E. J., 115 


326 


Egira simplex, 134 
Ehrlich, P. R., 91 
Eichlin, T. D., 193 
Elaphria festivoides, 312 
Ely, C. A., butterfly records from Kansas, 
256 
Enypia griseata, 131 
E. venata, 131 
Eomichla hallwachsae, 155 
Epargyreus antaeus, 170 
E. spanna, 170 
Erblichia odorata, 70 
Erynnis afranius, 92 
E. funeralis, 272 
Eucalyptus spp., as host plants, 91 
Eucosmomorpha albersana, 88 
Eucraera salambo, 91 
Eueides isabellae zoracaon, 321 
E. procula vulgiformis, 70 
Eueretagrotis perattenta, 312 
Eumeta rougeoti, 91 
Euphydryas gillettii, 92 
Eupithecia catalinata, 134 
Euproctis molunduana, 91 
Eupsilia morrisoni, 312 
E. tristigmata, 312 
E. vinulenta, 312 
Eurois astricta, 312 
Eurytides marcellus, 81 
Euthyatira pennsylvanica, 179 
E. pudens, 179 
Eutricopis nexilis, 20 
Euxoa divergens, 312 
E. longidentifera, 91 
E. perpolita, 312 
E. scandens, 312 
E. tristicula, 312 
Feltia heralis, 312 
Ferguson, D. C., 24, 96, 146, 179, 189 
Ferris, C. D., 187 
Finkelstein, I. L., 322 
Fixsenia ontario, 97 
Foeniculum vulgare, 29 
Forcipomyia fuliginosa, 185 
Gaddy, L. L., 166 
(Galli, 18, 164 Nas, Wal 
Galle iby 38.. ce, Wa 
Garnett, W. J., 289 
Gifford, S. M., 97 
Glaucopsyche lygdamus couperi, 258 
Gluck, W., butterfly records from Zaire, 
253 
Glycine max, 217 
Griselda radicana, 138 
Habeck, D. H., 224 
Hammond, P. C., 115 
Hardwick, D. F., 18, 148 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


Heliconius hecale zuleika, 70 
Heliothis armigera, 91 
Hemileuca maia, 168 
Herculia tenuis, 91 
Hesperia iowa, 181 
Hinchliff, J., 262 
Homorthodes furfurata, 306, 312 
H. lindseyi, 309 
H. uniformis, 306 
Iftner, D. C., 81 
Imelda mycea glaucomia, 255 
I. m. mycea, 255 
Immature Stages (descriptive): 
Ovay 22 lb2 95 
Larva, 22, 40, 74, 152) 183) V95ee2s: 
249, 306 
Pupa, 22, 41, 76, 158, 175, 183, 195, 218 
Incisalia henrici, 97 
Irwin, R. R., 260 
Janzen, D. H., 70 
Johnson, K., 78 
Juniperus virginiana, 78 
Kelson, R. V., 310 
Kern County, Calif. butterflies, 46 
Keuhn, R. M., new Wisconsin butterfly 
records, 228 
Knudson, E. C., 140 
Kotochalia junodi, 91 
Labedz, T. E., 256 
Lacanobia atlantica, 312 
L. grandis, 312 
L. legitima, 312 
L. lilacina, 312 
L. lutra, 312 
Lacinipolia lorea, 312 
L. olivacea, 312 
L. renigera, 312 
Latoia chapmanni, 91 
Lattin, J. D., 262 
Laurie, P., 166 
Lechriolipis nigrivenis, 91 
Leeuw, I., 87 
Lethe anthedon, 93 
L. eurydice, 93 
Leucania inermis, 312 
Libytheana bachmanii larvata, 121, 272 
Lirimiris meridionalis, 182 
Lithophane innominata, 312 
Lobobunaea phaedusa, 91 
Luxiaria curvivena, 91 
Megisto cymela, 176 
Melanchra adjuncta, 312 
M. assimilis, 312 
Melanolophia canadaria, 312 
Melanoplus islandicus, 267 
Melipotes indomita, 89 


VOLUME 37, NUMBER 4 


Metalepsis fishii, 312 
Miles, N. J., 207 
Miller, L. D., 69, 95, 317 
Miller, T. A., 174, 319 
Miller, W. E., 88 
Mimosa biuncifera, 203 

M. pigra, 193 
Minno, M. C., 310 
Mitoura gryneus, 289 

M. hesseli, (78), 97, 289 

M. johnsoni, 310 

M. spinetorum, 310 
Myscelia antholia, 164 
Nadiasa cuneata, 91 
Nathalis iole, 272 
Neck, R. W., 121, 269, 320 
Neil, K. A., 14, 249, 258, 306, 311 
Noctua pronuba, 169 
Nudaurelia conradsi, 91 

N. cytherea, 91 

N. dione, 91 

N. gueinzii, 91 

N. krucki, 91 
Nymphalis antiopa, 1, 275 
Obituaries: 37, 206, 260, 262, 322 
O'Brien, M. F., new butterfly distributions 

for New York State, 92 

Ochropleura plecta, 312 
Opler, P. A., 97 
Orgyia leucostigma plagiata, 312 

O. I. sablensis, 312 ; 
Orthonama obstipata, 91 
Orthosia hibisci, 312 

O. revicta, 312 
Pachypasa papyri, 91 

P. subfascia, 91 
Panthiades m-album, 272 
Papilio multicaudatus, 92 

P. rutulus, 275 

P. victorinus, 318 

P. zelicaon, 29 
Parasites of Lepidoptera: 239 
Parategeticula pollenifera, 207 
Parnassius clodius, 92 
Parsons, M., 83 
Passoa, S., 38, 193, 217 
Peridroma saucia, 312 
Pero morrisonaria, 312 
Phaneta argutipunctana, 143 

P. granulatana, 143 

P. linitipunctana, 140 
Phlagophora iris, 312 
Phoebis argante, 314 

P. sennae, 166, 272 
Pityopsis graminifolia, 149 
Plusia limbirena, 91 

P. putnami, 312 


327 


Polia imbrifera, 312 

P. latex, 312 
Polites coras, 267 

P. themistocles, 267 
Polygonia comma, 1 
Predators of Lepidoptera: 185, 242, 295 
Pseudaletia unipuncta, 312 
Pseudopanthera ennomosaria, 147 
Pseudorthodes vecors, 312 
Pseudothyatira cymatophoroides, 179 

P. expultrix, 179 
Pteryxia terebinthina, 34 
Rausher, M. D., 81 
Riotte, J. C. E., 89 
Rolfs, M. E., 256 
Rothschildia forbesi, 174 
Satyrium calanus falacer, 97 

S. kingi, 97 

S. liparops, 97 
Schwartz, A., 164 
Schweitzer, D. F., 85, 301 
Sevastopulo, D. G., 91 
Shapiro, A. M., 236, 259, 281 
Shawn KG? I 
Shinia honesta, 18 

S. rufipenna, 148 

S. tuberculum, 148 

S. verna, 18 
Sicyopsis blanchardata, 24 
Sideridis maryx, 249, 312 
Sims, S. R., 29, 286, 281 
Siproeta stelenes biplagiata, 320 
Smith: EV. 275 
Spartiniphaga carterae, 301 

S. inops, 301 
Speyeria atlantis elko, 244 

S. coronis, 92 

S. egleis moecki, 115 

S. e. oweni, 115 

S. e. tehachapina, 318 

S. nokomis, 92 
Spodoptera litteralis, 91 
Stamp, N. E., 145 
Stein, C., 129 
Stevens, R. E., 129 
Strymon melinus franki, 272 
Sylepta balteata, 91 
Synanthedon exitiosa, 87 
Syngrapha angulidens, 134 
Taraxacum officinale, 306 
Taylor, T. R., 160 
Tegeticula yuccasella, 207 
Theobroma cacao, 182 
Thymelicus lineola, 265 
Thyris maculata, 87 
Tmolus azia, 89 
Tricholita signata, 312 


328 


Trichoplusia ni, 160 
Troides oblongomaculatus papuensis, 83 
Urota sinope, 91 
Vanessa atalanta, 1 
V. a. rubria, 272 
V. cardui, 92 
West, D. A., 90 
Whittaker, P. L., satyrid populations in 
Costa Rica, 106 
Williams, T. S., 176 
Wright, B., 169 
Xestia dolosa, 312 
X. normaniana, 312 


JOURNAL OF THE LEPIDOPTERISTS SOCIETY 


X. oblata, 312 

X. smithii, 312 
Xylena curvimacula, 312 
Young, A. M., 186, 313 
Ypsolophus nella, 134 
Yucca baccata, 208 

Y. brevifolia, 208 

Y. elata, 208 

Y. torreyi, 208 

Y. whipplei, 208 
Zale minerea, 312 
Zeiraphera hesperiana, 138 
Zerene cesonia, 321 


a 


6 eT ee es Se a. — dem” 


EDITORIAL STAFF OF THE JOURNAL 
THoMaS D. EICHLIN, Editor 


% Insect Taxonomy Laboratory 
1220 N Street 
Sacramento, California 95814 U.S.A. 


Macpa R. Papp, Editorial Assistant 
DOUGLAS C. FERGUSON, Associate Editor THEODORE D. SARGENT, Associate Editor 
NOTICE TO CONTRIBUTORS 


Contributions to the Journal may deal with any aspect of the collection and study of 
Lepidoptera. Contributors should prepare manuscripts according to the following instruc- 
tions. 

Abstract: A brief abstract should precede the text of all articles. 

Text: Manuscripts should be submitted in triplicate, and must be typewritten, en- 
tirely double-spaced, employing wide margins, on one side only of white, 8% x 11 inch 
paper. Titles should be explicit and descriptive of the article’s content, including the 
family name of the subject, but must be kept as short as possible. The first mention of a 
plant or animal in the text should include the full scientific name, with authors of 
zoological names. Insect measurements should be given in metric units; times should be 
given in terms of the 24-hour clock (e.g. 0930, not 9:30 AM). Underline only where 
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SHEPPARD, P. M. 1959. Natural selection and heredity. 2nd. ed. Hutchinson, London. 
209 pp. 

196la. Some contributions to population genetics resulting from the study of 

the Lepidoptera. Adv. Genet. 10: 165-216. 


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CONTENTS 


CYPRIPEDIUM FLOWERS ENTRAP ADULT THYMELICUS (LEPI- 
DOPTERA: HESPERIIDAE) IN NORTHERN MICHIGAN. Ed- 
ward M: Barrows). i 


SIGNIFICANCE OF VISITS BY HACKBERRY BUTTERFLIES 
(NYMPHALIDAE: ASTEROCAMPA) TO FLOWERS. Raymond 
Wor Neck i ea a ee 


A TWELVE YEAR COUNT OF THREE CALIFORNIA BUTTER- 
FLIES. Leslie 'V> Smitth co a 


SEASONAL PHENOLOGY OF BATTUS PHILENOR (L.) (PAPILIONI- 
DAE) IN CALIFORNIA. S. R. Sims & A. M. Shapiro ....... 


DISTRIBUTION AND NOTES ON THE GREAT DISMAL SWAMP 
POPULATION OF MITOURA HESSELI RAWSON AND ZIEGLER 
(LYCAENIDAE). Andrew F. Beck & William J. Garnett .... 


A NEw SPECIES OF SPARTINIPHAGA (NOCTUIDAE) FROM THE NEW 
JERSEY PINE BARRENS. Dale F. Schweitzer 0000 


THE LARVA OF HOMORTHODES FURFURATA (GRT.) (NOG ra 
DAE)... Kenneth A. Neil) 


GENERAL NOTES 


Observations of hilltopping Mitoura spinetorum and M. johnsoni (Lycaeni- 
dae) in California. Richard V. Kelson ¢ Marc C. Minno 0c 


Lepidoptera reared on a simple wheat germ diet. Kenneth Neil WW. 


Egg placement by Phoebis (Pieridae) on Cassia (Leguminosae) “anticipates ’ 
the tropical rainy season, Allen M. Young 00 


Chermock, Hovanitz and Weber collections donated to Allyn Museum. Lee 
De Millen: 255 Ss eee in 


An old first United States record finally published: Papilio victorinus (Papi- 
lionidae) in Laredo, Texas. Jarmes K. AQ res oe.e...ccccccsccesssssssesceseeeecnsescneceserttmsneene 


Estimation of daily oviposition rates in reared female Antheraea polyphemus 
(Saturniidae). Thomas A. Miller Ge William J. COOg C8 ccccceccsecessessereeeee 


Additional comments on the butterflies of the Austin, Texas, region. Ray- 
morned Ws INGCK wie BA Oe Te ee 


CUBUT WAR TES a NCS 0 aE ee 


INDEX: TOs VOLUME Oe a : 


265 


269 


279 


281 


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